Vol 10 Part 14. An introduction to the immature stages of British Flies ...
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Royal En<strong>to</strong>mological Society<br />
HANDBOOKS FOR<br />
THE IDENTIFICATION<br />
OF BRITISH INSECTS<br />
To purchase current handbooks and <strong>to</strong> download<br />
out-<strong>of</strong>-print parts visit:<br />
http://www.royensoc.co.uk/publications/index.htm<br />
This work is licensed under a Creative Commons<br />
Attribution-NonCommercial-ShareAlike 2.0 UK:<br />
England & Wales License.<br />
Copyright © Royal En<strong>to</strong>mological Society 2013
Handbooks for <strong>the</strong> Identification <strong>of</strong> <strong>British</strong> Insects<br />
<strong>Vol</strong>. <strong>10</strong>, <strong>Part</strong> 14<br />
AN INTRODUCTION TO<br />
THE IMMATURE<br />
STAGES OF<br />
BRITISH FLIES<br />
DIPTERA LARVAE, WITH NOTES ON EGGS,<br />
PUP ARIA AND PUPAE<br />
K. G. V. Smith<br />
ROYAL ENTOMOLOGICAL SOCIETY OF LONDON
Handbooks for <strong>the</strong><br />
Identification <strong>of</strong> <strong>British</strong> Insects<br />
<strong>Vol</strong>. <strong>10</strong>, <strong>Part</strong> 14<br />
Edi<strong>to</strong>rs: W. R. Dolling & R. R. Askew<br />
AN INTRODUCTION<br />
TO THE<br />
IMMATURE STAGES<br />
OF<br />
BRITISH FLIES<br />
DIPTERA LARVAE,<br />
WITH NOTES ON EGGS, PUP ARIA AND PUPAE<br />
By<br />
K. G. V. SMITH<br />
Department <strong>of</strong> En<strong>to</strong>mology<br />
<strong>British</strong> Museum (Natural His<strong>to</strong>ry)<br />
London SW7 5BD<br />
1989<br />
ROYAL ENTOMOLOGICAL SOCIETY OF LONDON
The aim <strong>of</strong> <strong>the</strong> Handbooks is <strong>to</strong> provide illustrated identification keys <strong>to</strong> <strong>the</strong> insects <strong>of</strong> Britain,<br />
<strong>to</strong>ge<strong>the</strong>r with concise morphological, biological and distributional information.<br />
Each handbook should serve both as an <strong>introduction</strong> <strong>to</strong> a particular group <strong>of</strong> insects and as an<br />
identification manual.<br />
Details <strong>of</strong> handbooks currently available can be obtained from Publications Sales, <strong>British</strong><br />
Museum (Natural His<strong>to</strong>ry), Cromwell Road, London SW7 5BD.<br />
Cover illustration: egg <strong>of</strong> Muscidae; larva (lateral) <strong>of</strong> Lonchaea (Lonchaeidae); floating puparium<br />
<strong>of</strong> Elgiva rufa (Panzer) (Sciomyzidae).<br />
To Vera, my wife,<br />
with thanks for sharing<br />
my interest in insects<br />
World List abbreviation: Handbk /dent. Br./nsects.<br />
© Royal En<strong>to</strong>mological Society <strong>of</strong> London, 1989<br />
First published 1989 by <strong>the</strong> <strong>British</strong> Museum (Natural His<strong>to</strong>ry), Cromwell Road, London<br />
SW7 5BD.<br />
ISBN 0 901546 75 5<br />
Printed by Henry Ling Ltd., at <strong>the</strong> Dorset Press, Dorchester, Dorset.
Contents<br />
Page<br />
Introduction . 3<br />
Acknowledgements . 4<br />
Life-his<strong>to</strong>ries and ecology . 6<br />
Agricultural and economic importance 11<br />
Medical, environmental health and veterinary importance 15<br />
Preda<strong>to</strong>rs, parasites and pathogens. 19<br />
Fossil larvae and pupae 21<br />
Collecting, rearing, preservation and examination . 21<br />
Biology and morphology <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> 24<br />
Eggs. 24<br />
Larvae . 25<br />
~~ ~<br />
Classification, nomenclature and use <strong>of</strong> keys. 30<br />
Notes on <strong>the</strong> illustrations . 30<br />
Key <strong>to</strong> suborders <strong>of</strong> <strong>British</strong> Diptera larvae . 32<br />
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Nema<strong>to</strong>cera . 33<br />
Notes on families <strong>of</strong>Nema<strong>to</strong>cera 35<br />
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Brachycera . 55<br />
Notes on families <strong>of</strong> Brachycera . 57<br />
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Cyclorrhapha 71<br />
Notes on families <strong>of</strong> Cyclorrhapha 76<br />
Aschiza . 76<br />
Schizophora 85<br />
Acalyptratae. 85<br />
Calyptratae 115<br />
References. 141<br />
F~~- !M<br />
Index 259<br />
Introduction<br />
Knowledge <strong>of</strong> <strong>the</strong> larvae <strong>of</strong> Diptera lags far behind that <strong>of</strong> <strong>the</strong> adults. Of <strong>the</strong> 130 or<br />
so families <strong>of</strong> Diptera currently recognised in <strong>the</strong> world some 20 remain undescribed in<br />
<strong>the</strong> larval <strong>stages</strong>, including <strong>the</strong> following which occur in <strong>the</strong> <strong>British</strong> Isles: Acar<strong>to</strong>phthalmidae,<br />
Asteiidae, Camillidae, Chyromyidae, Stenomicridae, Tethinidae and Trixoscelidae<br />
(formerly included in <strong>the</strong> Heleomyzidae). These families are, however,<br />
included in <strong>the</strong> present handbook with suggestions as <strong>to</strong> where <strong>the</strong>ir larvae should be<br />
sought. Of <strong>the</strong> more than 80,000 species <strong>of</strong>Diptera known <strong>to</strong> science probably less than<br />
two per cent have been described in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> while adults <strong>of</strong> fur<strong>the</strong>r new<br />
species are continually being described.<br />
The families known best in <strong>the</strong> larval stage are usually those <strong>of</strong> medical or economic<br />
importance (e.g. Culicidae, Tipulidae). More recently families whose larvae are <strong>of</strong><br />
importance as ecological indica<strong>to</strong>rs are being systematically studied (e.g. Chironomidae).<br />
For <strong>the</strong> majority <strong>of</strong> families, however, only a few species come in<strong>to</strong> <strong>the</strong>se categories<br />
and description <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> has remained ra<strong>the</strong>r casual. Fortunately<br />
work on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> a particular family that occupies a distinct ecological<br />
niche has been recognised as a valuable research <strong>to</strong>pic for university students working<br />
3
for higher degrees. This has resulted in several valuable and comprehensive studies, e.g.<br />
Dixon (1960) and Hartley ( 1961) on Syrphidae; Okely ( 1974) and Pitkin ( 1988, in part)<br />
on Sphaeroceridae.<br />
Unfortunately all families do not lend <strong>the</strong>mselves <strong>to</strong> concentrated short term study<br />
and may require a long and continuous effort before a reasonably comprehensive<br />
treatment can be achieved, e.g. Brindle's work on Tipulidae (1952- 1967) and <strong>the</strong> 25<br />
years required <strong>to</strong> produce Skidmore's (1985) book on Muscidae. Some fine work has<br />
been done in Europe especially by Dusek & Laska (1967, summary) on Syrphidae, and<br />
Hennig (1943a-1956) and Schumann (1953- 1974) on Cyclorrhapha.<br />
The identification <strong>of</strong>larvae beyond family level has thus remained a difficult process<br />
requiring a knowledge <strong>of</strong>, and access <strong>to</strong>, a very widely scattered literature. In this<br />
Handbook keys <strong>to</strong> families and sub-families <strong>of</strong> final stage larvae are given as far as this is<br />
possible. In some families fur<strong>the</strong>r keys and illustrations are given <strong>to</strong> facilitate identification<br />
<strong>to</strong> genera and sometimes <strong>to</strong> species. For <strong>the</strong> latter <strong>the</strong> aim has been <strong>to</strong> include<br />
species <strong>of</strong> medical and economic importance, species commonly involved in <strong>the</strong> everyday<br />
enquiries <strong>of</strong> environmental health <strong>of</strong>ficers or <strong>the</strong> general public (including non<br />
<strong>British</strong> species regularly introduced in imported food, etc.) and those species most<br />
frequently encountered as larvae during ecological surveys or <strong>of</strong> special value in teaching<br />
biology at school or university level. To fur<strong>the</strong>r <strong>the</strong>se aims some bionomic keys are<br />
given at appropriate points where a limited number <strong>of</strong> species occur in a restricted<br />
habitat. In a section on ecology <strong>the</strong>re are some listings <strong>of</strong> families, genera or species <strong>to</strong><br />
be found in specific habitats which, by restricting <strong>the</strong> possibilities, should facilitate<br />
more rapid identification by reference <strong>to</strong> <strong>the</strong> figures alone. A full index provides rapid<br />
access <strong>to</strong> ecological information in <strong>the</strong> text. There are brief general comments on eggs<br />
and pupae, and illustrations showing <strong>the</strong>ir diversity <strong>of</strong> form.<br />
The following comprehensive specialist works on larvae are <strong>of</strong> value in identification.<br />
Hennig (1948- 1952) is well illustrated and lists <strong>the</strong> world species described as<br />
larvae up <strong>to</strong> that time with a full bibliography. Peterson (1957) treats Nearctic species<br />
but Teskey (in McAlpine et al., 1981) does so in much greater detail. Seguy (1950)<br />
gives much detailed information on all aspects <strong>of</strong> fly biology arranged under subject<br />
and habitat and similar information for <strong>British</strong> species is given in Stubbs & Chandler<br />
( 1978) which includes a key <strong>to</strong> families <strong>of</strong> larvae (by Brindle & Smith). Brauns ( 1954)<br />
deals with terrestrial Diptera larvae and pupae and is well illustrated. Teskey<br />
(1984) gives a well-illustrated key <strong>to</strong> aquatic Diptera larvae. Oldroyd & Smith (in<br />
Smith, 1973) give a key <strong>to</strong> families <strong>of</strong> larvae <strong>of</strong> medical importance. Smith ( 1986) deals<br />
with species <strong>of</strong> forensic importance including those found on carrion (human<br />
and animal). Koppen ( 1972) covers some agricultural pest species. Askew (1971) and<br />
Clausen (1940) review parasitic Diptera. References <strong>to</strong> particular families are given in<br />
each section below.<br />
As this book goes <strong>to</strong> press a very important two volume work has appeared on <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> Cyclorrhapha (Ferrar, 1987) and also part 2 <strong>of</strong> <strong>the</strong> Manual<br />
<strong>of</strong> Nearctic Diptera (McAipine et al., 1987), also covering Cyclorrhapha, has been<br />
published.<br />
Acknowledgements<br />
In his Introduc<strong>to</strong>ry Handbook <strong>to</strong> <strong>the</strong> Diptera <strong>the</strong> late Harold Oldroyd (1970b)<br />
expressed <strong>the</strong> hope that a volume dealing with <strong>the</strong> early <strong>stages</strong> <strong>of</strong> flies would eventually<br />
be produced in this series. Indeed, we had intended that we should cooperate in such a<br />
venture but his early death sadly prevented this. Never<strong>the</strong>less many <strong>of</strong> his ideas<br />
expressed both verbally and in his writings, especially in his unique book The Natural<br />
His<strong>to</strong>ry <strong>of</strong> <strong>Flies</strong> (1964), have stimulated an interest in <strong>the</strong> biology <strong>of</strong> flies among<br />
Dipterists, including <strong>the</strong> present author.<br />
4
The following colleagues working in <strong>the</strong> <strong>British</strong> Museum (Natural His<strong>to</strong>ry) (including<br />
those on <strong>the</strong> staff <strong>of</strong> <strong>the</strong> Commonwealth Institute <strong>of</strong> En<strong>to</strong>mology) are warmly<br />
thanked for reading appropriate sections <strong>of</strong> <strong>the</strong> work and for <strong>the</strong>ir constructive<br />
criticism, though <strong>the</strong>y must not <strong>of</strong> course be held responsible for any inaccuracies that<br />
remain: John Boorman, John E. Chainey, Dr Peter S. Crans<strong>to</strong>n, Dr Roger W.<br />
Crosskey, William R. Dolling, Dr Keith M . Harris (CIE), Dr Brian R. Pitkin, Adrian<br />
C. Pont, and Dr Ian M. White (CIE). Peter Chandler kindly checked <strong>the</strong> Myce<strong>to</strong>philidae<br />
and Platypezidae and Phi! Wi<strong>the</strong>rs checked Psychodidae.<br />
For generous permission <strong>to</strong> base some <strong>of</strong> my figures or keys on <strong>the</strong>ir published works<br />
(see References) I thank Dr E. J. Alien, Dr Pamela Alien, D . G. Ashby, Dr R. A .<br />
Bedding, Dr C. 0 . Berg, Dr P.A. Berry, Dr A. D. Bratt, Pr<strong>of</strong>. A. Brauns, Alan Brindle,<br />
Or D . E. Bryce, Or A. E. F. Chandler, Peter Chandler, Dr Lanna Cheng, Or W.<br />
Ciampolini, Dr C. P. Clausen, Or P. S. Crans<strong>to</strong>n, Or R. W. Crosskey, Dr R . Dahl, Or<br />
R. H . L. Disney, Or J . Doskocil, Pr<strong>of</strong>. C. Dupuis, Or A. Draber-Monko, Or J. Dusek,<br />
C. E. Dyte, Or W. C. Eastin, Dr H . J . Egglishaw, Or B. A. Foote, Or Paul Freeman,<br />
Or A. Friedberg, Pr<strong>of</strong>. P. Goeldlin, Pr<strong>of</strong>. W. L. Gojmerac, J . F. Graham, Dr D . J.<br />
Grea<strong>the</strong>ad, Dr L. R. Guima raes, Or K. M. Harris, Dr J. C. Hartley, Pr<strong>of</strong>. Wolfgang<br />
Hennig (for <strong>the</strong> late Pr<strong>of</strong>. Willi Hennig), Or D. F. Houlihan, Or C. Juliard, Or L. V.<br />
Knutson, Or N . P. Krivosheina, Or B. R. Laurence, Or A. Z . Lehrer, D . C. Lewis, Dr<br />
A. M . Lobanov, Or M. L. Luff, Dr L. Lyneborg, Dr M . E. Maldague, Or B. M.<br />
Mamaev, Or E. Mellini, Dr R . M . Miller, Or Hildegarde Miiller, Or A. Naga<strong>to</strong>mi, Or<br />
S. E. Neff, Or P. Nielsen, Or I. W. B. Nye, Pr<strong>of</strong>. T. Okada, P. Osborne, Dr V. Passovici,<br />
P. I. Persson, A. Peterson, Chih Ping, Or J . Robinson, G . H . R. Rothschild, R.<br />
Rozkosny, Or H. Schumann, Or J. R. Seeger, P. Sen, K. W . Simpson, P. Skidmore,<br />
Or Kathryn M. Sommerman, Or K. A. Spencer, Or J. W . Stephenson, G. C. Steyskal,<br />
Or V. N . Tanasijtshuk, Dr H . J . Teskey, R. J. Terry, Or L. Tascas, Or F. Vaillant,<br />
Pr<strong>of</strong>. J. B. Wallace, Dr Barbara M . Walshe, Dr F. Zumpt and Or J . Zuska.<br />
I thank <strong>the</strong> following publishers, journals and institutions for permission <strong>to</strong> base<br />
figures on illustrations appearing in <strong>the</strong>ir publications (deceased or untracea ble<br />
authors are cited in brackets, see References): Acta En<strong>to</strong>mologica Bohemoslavaca,<br />
Prague; Akademie-Verlag GmbH, Berlin (Hennig, Die Larvenformen der Dipteren);<br />
American En<strong>to</strong>mological Society, Philadelphia (Ent. News, Transactions); American<br />
Mosqui<strong>to</strong> Control Association, Washing<strong>to</strong>n DC (Mosqui<strong>to</strong> News); Association <strong>of</strong><br />
Applied Biologists, Wellesbourne (<strong>An</strong>n. appl. Bioi. , Barnes, Cohen, Steel, Thomas);<br />
Beitriige zur En<strong>to</strong>mologie, Berlin (Miiller); <strong>British</strong> En<strong>to</strong>mological and Natural His<strong>to</strong>ry<br />
Society, London (Proceedings); <strong>British</strong> Museum (Natural His<strong>to</strong>ry), London (Insects<br />
and O<strong>the</strong>r Arthropods <strong>of</strong> Medica/Importance, incl. drawings by <strong>the</strong> late Alan Palmer;<br />
Manual <strong>of</strong> Forensic En<strong>to</strong>mology); Butterworth Scientific Ltd., Guildford (Zumpt's<br />
Myiasis in Man and <strong>An</strong>imals in <strong>the</strong> Old World); Cahiers des Naturalistes, Paris (Cah.<br />
Na t.); Cambridge University Press (Parasi<strong>to</strong>logy, Bissett, Imms, Keilin, Rennie &<br />
Su<strong>the</strong>rland, Satchell, Saunders, Tate; J. mar. bioi. Ass. U.K., Sen); Czechoslovak<br />
Academy <strong>of</strong> Sciences in <strong>the</strong> publishing house Academia, Prague (Acta se. nat. Brno or<br />
Prirodov. Pr. Cesk. Akad. Ved.); C.A.B.International Institute <strong>of</strong> En<strong>to</strong>mology (Bull.<br />
ent. Res., Morris) Cornell University Agricultural Experimental Station, Ithaca;<br />
Dansk naturhis<strong>to</strong>risk Forening, Copenhagen; Deutsche En<strong>to</strong>mologische Zeitschrift,<br />
Berlin; Ejnar Munksgaard, Copenhagen and Reykjavik (Zoology <strong>of</strong> Iceland); En<strong>to</strong>mologica<br />
Scandinavica, Lund; En<strong>to</strong>mological Society <strong>of</strong> America, Maryland (<strong>An</strong>nals);<br />
En<strong>to</strong>mological Society <strong>of</strong> Washing<strong>to</strong>n, Washing<strong>to</strong>n DC (Proceedings); En<strong>to</strong>mologicheskoe<br />
Obozrenie, Moscow (Translation available as En<strong>to</strong>mological Review (Wiley),<br />
New York); En<strong>to</strong>mologiske Meddelelser, Copenhagen; En<strong>to</strong>mologist's Gazette (E. W.<br />
Classey Ltd.), Faringdon; En<strong>to</strong>mologist's Monthly Magazine (Gem Publishing Co.),<br />
Wallingford; Freshwater Biological Association, Ambleside (Scient. Pubis); Institut<br />
Royal des Sciences Naturelles de Belgique, Brussels (Bulletin); Dr W. Junk, The Hague,<br />
Bos<strong>to</strong>n, London (Series Ent.); Museum National d'His<strong>to</strong>ire Naturelle, Paris<br />
5
(Memoires); Museu de Zoologia, Universidad de Sao Paulo (Papeis Dep. Zoo/. S.<br />
Paulo); Parasitica, Gembloux (Seeger & Maldague); Royal En<strong>to</strong>mological Society <strong>of</strong><br />
London (Proceedings, Transactions, Handbk !dent. Br. Insects, <strong>An</strong>tenna, J . ent. (A) &<br />
(B), Syst. Ent.); Royal Society, London (Phi/. Trans. Hin<strong>to</strong>n, Mad war); Schweizerbart<br />
Verlag, Stuttgart (Die Fliegen der Palaearktischen Region, Engel, Hennig); Societe<br />
En<strong>to</strong>mologique Suisse, Lausanne (Mitt. Schweiz ent. Ges); Societe Zoologique de<br />
France, Paris (Bulletin); Staatlichen Museum fiir Naturkunde, Stuttgart (Stuttg. Beitr.<br />
Naturk.); Taylor & Francis Ltd., London (<strong>An</strong>n. Mag. nat. Hist., Keilin); Zoological<br />
Institute <strong>of</strong> <strong>the</strong> Polish Academy <strong>of</strong> Sciences, Warsaw; Institute <strong>of</strong> Evolutionary Morphology<br />
& Ecology <strong>of</strong> <strong>An</strong>imals, <strong>of</strong> <strong>the</strong> USSR Academy <strong>of</strong> Sciences and "Nauka"<br />
Publishers, Moscow; Zoological Society <strong>of</strong> London (Proceedings, Transactions, J.<br />
Zoo/., Frew); Zoologicheskii Zhurnal, Moscow.<br />
I thank <strong>the</strong> following for very kindly providing original pho<strong>to</strong>graphs: A. E. MeR.<br />
Pearce (figs 89-91), Dr K. M. Harris and J. Brown (figs 123-5) and B. van Aartsen and<br />
G. Helmers Jr. (fig. 270).<br />
For providing specimens, literature or information on Diptera larvae over <strong>the</strong> years I<br />
am indebted <strong>to</strong>: Dr Peter Bailey, L. Christie, Pr<strong>of</strong>. A. F . G . Dixon, Dr Y. Z.<br />
Erzinc;Iioglu, N. French, D. Green, Dr K. M . Harris, Pr<strong>of</strong>. B. Herting, <strong>the</strong> late Pr<strong>of</strong>. W.<br />
Hennig, <strong>the</strong> late Pr<strong>of</strong>. H. E. Hin<strong>to</strong>n, F.R.S., <strong>the</strong> late Dr B. M. Hobby, Dr A. G. Irwin,<br />
Dr D . P. Knight, Dr B. R. Laurence, Dr Jane Marshall, P. M. Miles, Dr A. J. Pontin,<br />
Dr M. W. Shaw, Dr M. C. D. Speight, C. Vardy, <strong>the</strong> late Pr<strong>of</strong>. G . C. Varley, P. Wi<strong>the</strong>rs,<br />
London and Whipsnade Zoos (Zoological Society <strong>of</strong> London), and all those veterinarians,<br />
pathologists, forensic scientists, environmental health and Ministry <strong>of</strong><br />
Agriculture (ADAS) <strong>of</strong>ficers <strong>to</strong>o numerous <strong>to</strong> mention here individually, though<br />
some are credited in <strong>the</strong> text for particular records.<br />
Adrian Pont very kindly helped me with translating <strong>the</strong> subtleties <strong>of</strong> <strong>the</strong> German and<br />
Russian languages, thus avoiding many misinterpretations from <strong>the</strong> literature.<br />
Finally I thank my wife for her careful and critical preparation <strong>of</strong> <strong>the</strong> typescript, her<br />
painstaking pro<strong>of</strong>-reading and compilation <strong>of</strong> <strong>the</strong> index, and her enthusiastic support<br />
throughout my en<strong>to</strong>mological activities.<br />
Life-his<strong>to</strong>ries and ecology<br />
<strong>Flies</strong> are highly adaptive insects and <strong>the</strong>ir larvae develop successfully in a very wide<br />
range <strong>of</strong> media, e.g. soil, rotting wood, plant and animal tissues (including man, alive or<br />
dead), fungi, dung and sewage, water (mostly fresh, sometimes brackish, occasionally<br />
marine), hot springs and geysers (some Ephydridae, Stratiomyidae) and even in petroleum<br />
pools (Halaeomyia, Ephydridae). Diptera exploit most <strong>of</strong> <strong>the</strong> available ecological<br />
niches in most <strong>of</strong> <strong>the</strong> biological roles, e.g. saprophages (many), commensals (some<br />
Sca<strong>to</strong>psidae, Chironomidae and Phoridae), preda<strong>to</strong>rs (many), parasites (many),<br />
symbionts (some Chironomidae and algae) and scavengers.<br />
Most <strong>of</strong> <strong>the</strong> protein feeding is done in <strong>the</strong> larval stage and <strong>the</strong> adults mostly need<br />
water <strong>to</strong> replace that lost during activity, especially flight, and sugars, from nectar or<br />
honeydew, <strong>to</strong> provide <strong>the</strong>ir energy. Some adults also take protein, e.g. preda<strong>to</strong>rs, blood<br />
feeders, pollen feeders.<br />
As a rule, aquatic larvae are carnivorous or live on algae. Apart from preda<strong>to</strong>rs,<br />
<strong>the</strong> precise nature <strong>of</strong> <strong>the</strong> food taken by terrestrial larvae is <strong>of</strong>ten inadequately<br />
known. Thus <strong>the</strong> larva <strong>of</strong> <strong>the</strong> bluebottle (Calliphora) is classed as a carrion feeder but in<br />
fact bacteria on <strong>the</strong> carrion are probably <strong>the</strong> main food source (Roberts, 1971 a).<br />
Similarly bacteria are probably <strong>the</strong> main source <strong>of</strong> food and vitamins for housefly<br />
larvae (Levinson, 1960). Baumberger( 1919) found that many supposedly saprophagous<br />
fly larvae are actually myce<strong>to</strong>phagous. Clearly <strong>the</strong>re is plenty <strong>of</strong> scope for careful<br />
research here.<br />
6
The ancestral type <strong>of</strong> habitat for Diptera is probably in bogs and swamps and among<br />
mosses and decaying wood on drier land. Thus aquatic and terrestrial types <strong>of</strong> larvae<br />
developed <strong>to</strong>ge<strong>the</strong>r and both are present in many families <strong>of</strong> Nema<strong>to</strong>cera (e.g.<br />
Tipulidae, Cera<strong>to</strong>pogonidae) and Brachycera (Stratiomyidae, Dolichopodidae, etc.).<br />
Many more Nema<strong>to</strong>cera than Brachycera became purely aquatic, <strong>the</strong> Brachycera tending<br />
<strong>to</strong>wards a more terrestrial mode <strong>of</strong> life. The larvae <strong>of</strong>Nema<strong>to</strong>cera and Brachycera<br />
<strong>of</strong>ten exhibit a wide range <strong>of</strong> adaptative forms, but in <strong>the</strong> higher Brachycera (e.g.<br />
terrestrial Empididae and Dolichopodidae) <strong>the</strong> larvae are more streamlined and nearer<br />
<strong>the</strong> maggot type <strong>of</strong> larvae typical <strong>of</strong> <strong>the</strong> Cyclorrhapha, though <strong>the</strong> Syrphidae<br />
(Cyclorrhapha Aschiza) also have a remarkably wide range <strong>of</strong> forms.<br />
Among <strong>the</strong> Cyclorrhapha, <strong>the</strong> Acalyptratae and Calyptratae have exploited most <strong>of</strong><br />
<strong>the</strong> possible habitats arising from <strong>the</strong> decomposing 'compost' type <strong>of</strong> breeding medium<br />
through dung and carrion <strong>to</strong> living plant and animal tissues.<br />
Some specific life his<strong>to</strong>ries are described under <strong>the</strong> Medical and Economic sections<br />
and under each family. Some specific ecological associations are listed here under <strong>the</strong><br />
sections on agricultural importance and, for Diptera occupying macro- and microhabitats,<br />
Stubbs & Chandler (1978), which complements Colyer & Hammond (1968),<br />
should be consulted. Seguy ( 1950) is also useful. Broader ecological issues are discussed<br />
by Oldroyd ( 1964). Some specific habitats occupied by Diptera in <strong>the</strong>ir larval <strong>stages</strong> are<br />
listed below, with an indication <strong>of</strong> those most likely <strong>to</strong> be found <strong>the</strong>re.<br />
Soil. Many families (except those with purely aquatic larvae) have some members<br />
which may occasionally be found in soil in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>, usually for <strong>the</strong> purpose<br />
<strong>of</strong> pupation. However <strong>the</strong> families most likely <strong>to</strong> be encountered in sampling soil<br />
are: Tipulidae, Bibionidae, Chironomidae, Therevidae, Stratiomyidae, Tabanidae,<br />
Rhagionidae, Asilidae, Empididae, Dolichopodidae, Phoridae, Sphaeroceridae and<br />
Muscidae.<br />
Dead wood. Over 200 species representing 42 families spend <strong>the</strong>ir <strong>immature</strong> <strong>stages</strong><br />
in dead wood. The type <strong>of</strong> wood can influence <strong>the</strong> fauna, conifers supporting<br />
different species <strong>to</strong> deciduous trees. Rates <strong>of</strong> decay between tree species also vary.<br />
However, since dead wood is not readily identifiable in practice, <strong>the</strong> records are<br />
grouped here.<br />
Keys <strong>to</strong> <strong>the</strong> larvae <strong>of</strong> wood-inhabiting Diptera are provided by Krivosheina &<br />
Mamaev (1967) and Mamaev, Krivosheina & Po<strong>to</strong>tskaya (1977) key species predaceous<br />
upon tree trunk 'pests'. Teskey ( 1976) deals with Nearctic species associated with<br />
trees. Kitching ( 1971) deals specifically with <strong>the</strong> fauna <strong>of</strong> rot-holes.<br />
Diptera most frequently encountered are Tipulidae (especially Ctenophora),<br />
Myce<strong>to</strong>philidae, Sciaridae, Cera<strong>to</strong>pogonidae (especially Forcipomyia), Cecidomyiidae<br />
(especially Lestremiinae), Stratiomyidae, Xylomyidae, Xylophagidae, Empididae (especially<br />
Tachydromiinae, Ocydromiinae), Dolichopodidae, Syrphidae, Pallopteridae,<br />
Lonchaeidae, Clusiidae and Muscidae (especially Phaonia). Where bark beetles<br />
(Sco/ytus) are present, larvae <strong>of</strong> Medetera (Dolichopodidae) and Odinia (Odiniidae)<br />
may occur. In nests <strong>of</strong> Sphecidae (Hymenoptera) <strong>the</strong> larvae or puparia <strong>of</strong> Sarcophagidae<br />
or Eusta/omyia (<strong>An</strong>thomyiidae) may be found.<br />
Puparia <strong>of</strong> parasitic species may also be found: Tachinidae, e.g. Triarthria (on<br />
earwigs), Loewia (on centipedes), Admontia (Trichopareia) (on tipulid larvae);<br />
Rhinophoridae (on woodlice).<br />
Dung. Skidmore (in Stubbs & Chandler, 1978) lists <strong>the</strong> <strong>British</strong> species associated with<br />
various types <strong>of</strong> dung and notes that about five per cent <strong>of</strong> <strong>the</strong> <strong>British</strong> Diptera (c. 300<br />
species) have been recorded as coprophilous, two per cent (c. 120 species) exclusively in<br />
<strong>the</strong> larval stage. Kumar & Lloyd (1976) provide a bibliography <strong>of</strong> arthropods associated<br />
with dung. The association <strong>of</strong> flies with dung has received considerable atten~ion<br />
7
due <strong>to</strong> <strong>the</strong> possible health hazards <strong>of</strong> adults flying from excrement <strong>to</strong> foodstuffs but<br />
precise ecological studies are few.<br />
The following papers deal with <strong>the</strong> fly faunas <strong>of</strong> different types <strong>of</strong> dung: cow<br />
(Laurence 1953b-55, Papp & Garz6, 1985); dog (Disney, 1973a, 1976a); human<br />
(Howard, 1900, Steyskal, 1957); horse and donkey (Papp, 1976); sheep (Papp, 1985).<br />
Of <strong>the</strong> 38 families listed by Skidmore <strong>the</strong> following are most frequently encountered<br />
as larvae in dung and listed in approximately <strong>the</strong> order <strong>of</strong> abundance found in cow<br />
dung by Laurence (1954): Psychodidae, Sphaeroceridae, Sepsidae, Scathophagidae,<br />
Trichoceridae, <strong>An</strong>isopodidae, Cera<strong>to</strong>pogonidae, Muscidae (many predaceous),<br />
Chironomidae (Smittia), Stratiomyidae, Sca<strong>to</strong>psidae, Empididae ( all predaceous),<br />
Cecidomyiidae, Syrphidae. The entry under dung in <strong>the</strong> index indicates references in<br />
<strong>the</strong> text <strong>to</strong> that <strong>of</strong> specific animals.<br />
Diptera breeding in sewage include: Psychodidae, Chironomidae, <strong>An</strong>isopodidae,<br />
Syrphidae (Eristalis), Ephydridae, Sphaeroceridae and Scathophagidae (Lloyd et al.,<br />
1940; Green, 1970; Busvine, 1980).<br />
Carrion. Following <strong>the</strong> death <strong>of</strong> an animal various insects and o<strong>the</strong>r invertebrates,<br />
but especially Diptera and Coleoptera, invade <strong>the</strong> corpse in a succession <strong>of</strong> about five<br />
distinct 'waves' <strong>of</strong> species. Each wave coincides with a particular stage <strong>of</strong> decay <strong>of</strong> <strong>the</strong><br />
corpse and may contain not only carrion feeders but also <strong>the</strong>ir preda<strong>to</strong>rs and parasites.<br />
The feeding <strong>of</strong> Diptera larvae rapidly accelerates <strong>the</strong> decomposition <strong>of</strong> <strong>the</strong> corpse.<br />
A general account <strong>of</strong> carrion fauna and succession (especially on man) is given in<br />
Smith (1986b) (see also under Medical Importance- Forensic). Detailed individual<br />
studies <strong>of</strong> succession involving Diptera on particular animals in Britain are few : snails<br />
(Beaver, R. A., 1971), rabbit (Chapman, R. F. & Sankey, 1955), voles (Lane, 1975),<br />
fox (Smith, 1975). The families important as larval carrion feeders are as follows<br />
in <strong>the</strong>ir approximate order <strong>of</strong> appearance on a corpse: Calliphoridae (Calliphora,<br />
Lucilia and later Cynomya, Phormia, Pro<strong>to</strong>phormia), Muscidae (Musca, Muscina),<br />
Sarcophagidae, Piophilidae, Fanniidae, Drosophilidae, Sepsidae, Syrphidae (Eristalis),<br />
Ephydridae (Teichomyza), Muscidae (Ophyra), Phoridae, Piophilidae (including<br />
Thyreophoridae).<br />
Water. The distinction between truly terrestrial and truly aquatic larvae is not always<br />
a clear one. Habitats range from marshy soil, among aquatic mosses, on rock face<br />
seepages, ditches, plant axils and tree rot holes <strong>to</strong> larger bodies <strong>of</strong> standing or flowing<br />
water.<br />
Most families <strong>of</strong> Nema<strong>to</strong>cera, several families <strong>of</strong> Brachycera and a few families <strong>of</strong><br />
Cyclorrhapha have truly aquatic <strong>immature</strong> <strong>stages</strong>. Exclusively aquatic: Chaoboridae,<br />
Culicidae, Dixidae, Ptychopteridae, Simuliidae and Thaumaleidae. Mostly aquatic:<br />
Cera<strong>to</strong>pogonidae, Chironomidae. Families with some aquatic members: Psychodidae,<br />
Tipulidae, Stratiomyidae (Stratiomyinae, Clitellariinae), Rhagionidae (A<strong>the</strong>rix,<br />
Atrichops, now regarded as a separate family A<strong>the</strong>ricidae), Tabanidae (mostly wet soil,<br />
Chrysops and some Hybomitra truly aquatic), Empididae (Hemerodromiinae, Clinocerinae,<br />
Oreoge<strong>to</strong>ninae), Dolichopodidae, Syrphidae (Eristalini and Chrysogaster),<br />
Ephydridae, Sciomyzidae, Scathophagidae (Spaziphora, Hydromyza), Muscidae (most<br />
Limnophorinae, probably some Coenosiinae, Phaonia exoleta Meigen in tree rot holes<br />
(see also under plants), Graphomya maculata Scopoli in mud in puddles).<br />
The few marine species are dealt with in <strong>the</strong> next section.<br />
Sea shore. Diptera <strong>of</strong> marine environments are treated in Cheng ( 1976). Some breed<br />
in <strong>the</strong> lit<strong>to</strong>ral or intertidal zone: Chironomidae (Thalassomya, Psamathiomya, Halocladius,<br />
Clunio and Thalassosmittia), Tipulidae (Limonia (Geranomyia) unicolor<br />
Haliday), Dolichopodidae (Aphrosylus, Dolichopus nubilis Meigen, Machaerium ,<br />
Hydrophorus oceanus Macquart, Hygrocoleuthus, Rhaphium consobrinum Zetterstedt),<br />
8
Empididae ( Chersodromia-<strong>immature</strong> <strong>stages</strong> unknown), Syrphidae (Eristalinus<br />
aeneus (Scopoli)), Ephydridae (Parydroptera), Canacidae.<br />
On <strong>the</strong> drift line, among seaweed (see Backlund, 1945a), larvae <strong>of</strong> <strong>the</strong> following<br />
families occur: Coelopidae, Helcomyzidae, Sepsidae (Orygma), Sphaeroceridae<br />
(Thoracochaeta spp., Lep<strong>to</strong>cera (Rachispoda)fuscipennis Haliday), Scathophagidae<br />
(Scathophaga li<strong>to</strong>rea Fallen, S. calida Curtis, Ceratinos<strong>to</strong>ma ostiorum Curtis), <strong>An</strong>thomyiidae<br />
(Fucellia), Fanniidae (Fannia canicularis).<br />
Coastal sand dunes may be a harsh environment but Ardi:i (1957) found 750 species<br />
<strong>of</strong> Diptera on dunes in Sweden. However, less than 50 species were confined <strong>to</strong> dunes<br />
(steno<strong>to</strong>pic). The following Diptera are known <strong>to</strong> breed in <strong>the</strong> sand: Sca<strong>to</strong>psidae<br />
(Aspistes berolinensis Meigen), Tipulidae (Tipulajuncea Meigen), Asilidae (Philonicus<br />
a/biceps Meigen), Therevidae (Thereva annulata F.), Chamaemyiidae (larvae predaceous<br />
on coccids at base or roots <strong>of</strong> grasses), Trixoscelidae (Trixoscelis puparia have<br />
been found in sand under sea sandwort Honkenya, but <strong>the</strong> larva is undescribed and<br />
probably develops in guano), <strong>An</strong>thomyiidae (Delia spp., probably on roots <strong>of</strong> grasses).<br />
In view <strong>of</strong> <strong>the</strong> number <strong>of</strong> species associated with this habitat, and <strong>the</strong> few species<br />
reared, <strong>the</strong>re is clearly scope for fur<strong>the</strong>r research.<br />
Preda<strong>to</strong>rs. Many truly aquatic Nema<strong>to</strong>cera larvae, many terrestrial Brachycera, but<br />
only a few Cyclorrhapha are predaceous in <strong>the</strong> larval stage. The prey usually consists <strong>of</strong><br />
o<strong>the</strong>r insect larvae or small worms. Some families are specialist feeders, e.g. Syrphini on<br />
aphids, Sciomyzidae on molluscs. The family distribution <strong>of</strong> <strong>the</strong> predaceous habit in<br />
larvae is as follows: Tipulidae (Pediciini, Hexa<strong>to</strong>mini), Myce<strong>to</strong>philidae (Keroplatiinae),<br />
Cecidomyiidae (some Cecidomyiidi), Chaoboridae, Culicidae (only Toxorhynchitini),<br />
Cera<strong>to</strong>pogonidae (Cera<strong>to</strong>pogoninae), Chironomidae (Tanypodinae, some Chironominae,<br />
few Orthocladinae), Tabanidae, Rhagionidae (including A<strong>the</strong>ricidae), Xylophagidae,<br />
Xylomyidae, Stratiomyidae (Pachygastrinae), Therevidae, Scenopinidae,<br />
Asilidae, Empididae, Dolichopodidae, Phoridae, Syrphidae (on aphids, etc.), Chamaemyiidae<br />
(on Homoptera), Drosophilidae (Acle<strong>to</strong>xenus on Homoptera), Chloropidae<br />
( Chloropisca on sugar beet root aphid), Sciomyzidae (molluscs), Scathophagidae,<br />
\-fuscidae (Limnophorinae, Coenosiinae, Phaoniinae some Mydaeinae, Muscina,<br />
Ophyra, Hydrotaea, etc.), <strong>An</strong>thomyiidae (some Delia, etc.).<br />
Parasites and parasi<strong>to</strong>ids. Parasitic larvae are infrequent in <strong>the</strong> Nema<strong>to</strong>cera and<br />
Brachycera and become more common in <strong>the</strong> Cyclorrhapha where it is <strong>the</strong> normal<br />
lifestyle for some families, e.g. Conopidae, Tachinidae. Some families have parasitic<br />
5pecies abroad but not in Britain and o<strong>the</strong>rs have a wider range <strong>of</strong> hosts abroad, e.g.<br />
Bombyliidae, Conopidae. The parasitic habit is distributed in <strong>British</strong> families as follows.<br />
The hosts in Britain are indicated in brackets (see also under hosts). Useful<br />
references are Clausen (1940), Askew (1971): Chironomidae (Parachironomus,<br />
G(rp<strong>to</strong>tendipes in aquatic molluscs), Cecidomyiidae (Homoptera), Acroceridae<br />
(spiders), Bombyliidae (Hymenoptera, Orthoptera, Lepidoptera), Phoridae (Phalacro<strong>to</strong>phora<br />
on Coccinellidae (Coleoptera), Borophaga incrassata Meigen on Bibio<br />
IDiptera), Pseudacteon on ants), Pipunculidae (Homoptera), Chloropidae (Siphonella<br />
in spider egg sacs), Sciomyzidae (parasi<strong>to</strong>id on molluscs), Calliphoridae (Pollenia and<br />
Be/lardia in earthworms; Melinda, Eggisops, molluscs), Rhinophoridae (woodlice),<br />
Sarcophagidae (Senotainia, wasps; Sarcophaga, spider egg cocoons, insects, etc, snails),<br />
Oestridae (sheep, deer), Hypodermatidae (cattle, horses, deer), Gasterophilidae<br />
( bo~). Tachinidae (insects, including o<strong>the</strong>r Diptera).<br />
Nests- The larvae <strong>of</strong> several families are specifically associated with nests <strong>of</strong> social<br />
insects, birds or mammals, as commensals, scavengers or parasites as follows.<br />
Nests <strong>of</strong> social insects (useful reference O'Toole in Stubbs & Chandler, 1978):<br />
Cera<strong>to</strong>pogonidae, Sciaridae, Sca<strong>to</strong>psidae (ants), Bombyliidae (bees), Phoridae (e.g.<br />
9
Aenigmatias on ants), Syrphidae (Microdon, ants; <strong>Vol</strong>ucella, wasps), Sphaeroceridae,<br />
Milichiidae (ants), Braulidae (honey-bees), Sarcophagidae (Mil<strong>to</strong>gramminae, wasps<br />
and bees), Fanniidae, Muscidae.<br />
Birds' nests (useful references: Hicks, 1959, 1963a, 1971; Woodr<strong>of</strong>fe, 1953): Sca<strong>to</strong>psidae,<br />
Chironomidae (Bryophaenocladius nitidico/lis (Goetghebuer)), Scenopinidae,<br />
Phoridae, Chyromyidae, Piophilidae (Neottiophilinae), Carnidae, Heleomyzidae,<br />
Trixoscelidae, Sphaeroceridae, Fanniidae, Muscidae (some Helina, Hydrotaea, Muscina,<br />
Ophyra), <strong>An</strong>thomyiidae (<strong>An</strong>thomyia), Sarcophagidae, Calliphoridae (especially<br />
Pro<strong>to</strong>ca/liphora, Calliphora), Hippoboscidae.<br />
Mammal burrows and nests (useful references: Hackman, 1967; Hutson, in<br />
Stubbs & Chandler, 1978): Heleomyzidae, Sphaeroceridae, Phoridae, Camillidae,<br />
Sciaridae.<br />
Fuller lists <strong>of</strong> rea rings <strong>of</strong>Diptera from nests are given in <strong>the</strong> references cited. Careful<br />
work is still needed on <strong>the</strong> ecology <strong>of</strong> nest-frequenting Diptera. E. B. Basden's extensive<br />
collection <strong>of</strong>Diptera from nests is available for study in <strong>the</strong> Royal Museum <strong>of</strong> Scotland<br />
(see Ro<strong>the</strong>ray, 1989 for summary).<br />
Plant feeders. The larval <strong>stages</strong> <strong>of</strong> many Diptera are described in <strong>the</strong> literature as<br />
living in and feeding on 'decaying vegetation'-a category not specifically included in<br />
this Handbook, except for dead wood. However, many terrestrial (some aquatic larvae<br />
feed on algae and mosses) species feed on <strong>the</strong> living tissues <strong>of</strong> plants in specific ways as<br />
listed below. Records marked with a need fur<strong>the</strong>r investigation or confirmation.<br />
Larvae <strong>of</strong> agricultural importance are treated more fully in <strong>the</strong> section so titled.<br />
Surface leaf feeders: Tipulidae (Cylindro<strong>to</strong>minae, mosses), Myce<strong>to</strong>philidae (Boletina,<br />
liverworts; Gnoriste, mosses; Docosia, lichens), Rhagionidae (Ptiolina on mosses).<br />
Leaf-miners: Cecidomyiidae (some Lestremiinae, liverworts; higher plants),<br />
Chironomidae, Sciaridae (Phy<strong>to</strong>sciara on Ranunculus), Rhagionidae (Spania nigra<br />
Meigen, liverworts), Dolichopodidae (Thrypticus, in stems), Syrphidae (some Cheilosia),<br />
Agromyzidae, Tephritidae (some), <strong>An</strong>thomyzidae (<strong>An</strong>agnota, Paranthomyza),<br />
Lauxaniidae, Psilidae, Drosophilidae (Scap<strong>to</strong>myza), Ephydridae, Scathophagidae,<br />
<strong>An</strong>thomyiidae (especially Pegomya). Mines caused by Diptera (unlike o<strong>the</strong>r orders)<br />
show primary and secondary feeding tracks causing a characteristic herring-bone pattern<br />
and linear mines have <strong>the</strong> frass lying alternately on <strong>the</strong> two sides (Hering, 1951);<br />
this is because mining Diptera larvae feed on <strong>the</strong>ir sides, facing first one direction, <strong>the</strong>n<br />
<strong>the</strong> o<strong>the</strong>r.<br />
Gall-formers: Cecidomyiidae, Tephritidae, Lauxaniidae (Calliopum, clover and<br />
Viola), Chloropidae (Lipara, reeds; Chlorops pumilionis Bjerkander, barley).<br />
Many o<strong>the</strong>r Diptera larvae feed in stems, roots, flowers, seeds and fruits, some <strong>of</strong><br />
which are <strong>of</strong> economic importance (see section on Agricultural Importance). Uffen &<br />
Chandler (in Stubbs & Chandler, 1978) list <strong>the</strong> Diptera associated with <strong>the</strong> higher<br />
plants. Teskey (1976) gives an account <strong>of</strong> Diptera larvae associated with trees (in<br />
North America), including those developing in sap exudates. Kitching (1971) deals<br />
specifically with <strong>the</strong> insect fauna <strong>of</strong> tree rot-holes (see index also).<br />
Fungi. The larval Diptera found in fungi have been <strong>the</strong> subject <strong>of</strong> some detailed<br />
studies (Bux<strong>to</strong>n, 1960; Smith, 1956; Trifourkis, 1977). Chandler (in Stubbs &<br />
Chandler, 1978) lists in detail <strong>the</strong> known fungus associations <strong>of</strong> <strong>British</strong> Diptera, which<br />
involve larvae <strong>of</strong> <strong>the</strong> following families: Trichoceridae, Tipulidae, Psychodidae, Cera<strong>to</strong>pogonidae,<br />
Chironomidae, <strong>An</strong>isopodidae, Myce<strong>to</strong>philidae, Sciaridae, Sca<strong>to</strong>psidae,<br />
Cecidomyiidae, Scenopinidae, Empididae (as preda<strong>to</strong>rs), Dolichopodidae, Phoridae,<br />
Platypezidae, Syrphidae (some Cheilosia), Platys<strong>to</strong>matidae, Dryomyzidae, Heleomyzidae,<br />
Sepsidae, Lauxaniidae, Sphaeroceridae, Lonchaeidae, Piophilidae, Odiniidae,<br />
Asteiidae, Drosophilidae, Chloropidae, Tachinidae (as parasites <strong>of</strong> Lepidoptera<br />
(Tineidae)), <strong>An</strong>thomyiidae, Fanniidae, Muscidae (as preda<strong>to</strong>rs).<br />
<strong>10</strong>
Only six cases <strong>of</strong>Diptera inducing galls in fungi are known: Myocecis ova/is Edwards<br />
(Cecidomyiidae) on Hypochnus; Agathomyia wankowiczi (Schnabl) (Platypezidae) on<br />
Ganoderma applanatum (Pers.) Pat.; Drosophila phalerata Meigen (Drosophilidae) on<br />
Psathyra; Myce<strong>to</strong>phila blanda Winnertz (Myce<strong>to</strong>philidae) on Lactarius deliciosus L. ex<br />
Fries and an undescribed dipteran on Daedalea quercina L. and Conocybe (Harris &<br />
Evans, 1979).<br />
Families recorded only as visiting adults are not included.<br />
Caves. Speolepta lep<strong>to</strong>gaster (Winnertz) (Myce<strong>to</strong>philidae) is <strong>the</strong> only <strong>British</strong> fly that<br />
is almost an exclusive cave-dweller (troglobite, troglobiont, troglobie). This species<br />
breeds in caves, <strong>the</strong> larva living in a slime tube attached by lateral strands <strong>to</strong> <strong>the</strong> damp<br />
cave walls and ceiling. The larva forages from <strong>the</strong> tube <strong>to</strong> feed on algae and fungi.<br />
The head capsule is pale, which gives it <strong>the</strong> superficial appearance <strong>of</strong> a nema<strong>to</strong>de.<br />
Myce<strong>to</strong>philidae <strong>of</strong> <strong>the</strong> genus Macrocera may also breed in caves.<br />
Frequent cave dwellers that also occur outside caves (troglophiles) are Trichocera<br />
maculipennis Meigen (Trichoceridae), Triphleba antricola (Schmitz) (Phoridae), and<br />
Terrilimosina racovitzai (Bezzi) (Sphaeroceridae).<br />
Heleomyza serrata (L.) and Scoliocentra villosa (Meigen) (Heleomyzidae) occur<br />
frequently in caves but are equally common outside. Chyromya species (Chyromyidae)<br />
also occur in caves. The dung <strong>of</strong> bats (or o<strong>the</strong>r mammals), fungi, o<strong>the</strong>r vegetation, or<br />
<strong>the</strong> presence <strong>of</strong> water may all serve as attractive breeding media <strong>to</strong> Diptera. The phorid<br />
Megaselia melanocephala parasitizes eggs <strong>of</strong> <strong>the</strong> common cave spider M eta menardi<br />
Latreille.<br />
Many adult Diptera may shelter or hibernate in caves (see Hutson in Stubbs &<br />
Chandler, 1978). M a tile (1970) reviews <strong>the</strong> Diptera fauna <strong>of</strong> caves worldwide.<br />
Agricultural and economic importance<br />
The larvae <strong>of</strong> a few Diptera regularly attack food crops and farm animals or important<br />
horticultural plants and may be regarded as pests. These are listed below and <strong>the</strong><br />
more significant species are also included in <strong>the</strong> notes, illustrations and some keys<br />
under <strong>the</strong> appropriate family. Occasional pest species are mentioned under <strong>the</strong><br />
appropriate family.<br />
The common names used follow Seymour (1979) and useful general texts are<br />
Edwards, C. A. & Heath (1964) and Buczacki & Harris (1981).<br />
TIPULIDAE<br />
Tipula o/eracea Linnaeus<br />
(crane fly, lea<strong>the</strong>rjacket)<br />
• T. pa/udosa Meigen<br />
(crane fly, lea<strong>the</strong>rjacket)<br />
Nephro<strong>to</strong>ma appendiculata (Pierre)<br />
(crane fly, lea<strong>the</strong>rjacket)<br />
Oats, wheat, turnip, mangold, pota<strong>to</strong>. Worst<br />
damage in spring, especially on newly ploughed<br />
land<br />
Dit<strong>to</strong><br />
Dit<strong>to</strong>, more frequently in gardens<br />
BIBIONIDAE<br />
Bibio marci (Linnaeus)<br />
(St. Mark's fly)<br />
Bibio, o<strong>the</strong>r spp.<br />
(March flies)<br />
Dilophusfebrilis (Linnaeus)<br />
(fever fly)<br />
CHIRONOMIDAE<br />
Bryophaenocladius<br />
Smillia<br />
Celery, lettuce, asparagus, grass, Polyanthus.<br />
Sax ifraga<br />
Winter wheat, grass, celery, sugarbeet, etc. (see<br />
Freeman & Lane, 1985)<br />
Spring barley, grass, maize, sugarbeet, pota<strong>to</strong>,<br />
lettuce, hops, <strong>to</strong>ma<strong>to</strong>, chrysan<strong>the</strong>mum<br />
Roots <strong>of</strong> greenhouse plants<br />
Barley, winter corn, winter wheat<br />
11
SCIARIDAE<br />
* Bradysia brunnipes (Meigen)<br />
(mushroom Sciarid)<br />
* Lycoriella auripila (Winnertz)<br />
(mushroom Sciarid)<br />
* L. so/ani (Winnertz)<br />
(mushroom Sciarid)<br />
Plas<strong>to</strong>sciara perniciosa (Edwards)<br />
Pnyxia scabei (Hopkins)<br />
CECIDOMYIIDAE<br />
Campylomyza ormerodi (Kieffer)<br />
(red clover gall gnat)<br />
Contarinia humuli (Theobald)<br />
(hop strig maggot)<br />
C. nasturtii (Kieffer)<br />
(swede midge)<br />
C. pisi (Winnertz)<br />
(pea midge)<br />
C. pyrivora (Riley)<br />
(pear midge)<br />
C. tritici (Kirby)<br />
(wheat midge)<br />
Dasineura brassicae (Winnertz)<br />
(brassica pod midge)<br />
D. leguminicola (Lintner)<br />
(clover seed midge)<br />
D. tetensi (Riibsaamen)<br />
(blackcurrant leaf midge)<br />
D. trifolii (Low, F.)<br />
(clover leaf midge)<br />
Halodiplosis marginata (V on Roser)<br />
(saddle gall midge)<br />
Henria psalliotae Wyatt<br />
(mushroom cecid)<br />
Heteropeza pygmaea Winnertz<br />
(mushroom cecid)<br />
Lestremia cinerea Macquart<br />
(mushroom midge)<br />
Mycophila barnesi Edwards<br />
(mushroom cecid/midge)<br />
M. speyeri (Barnes)<br />
(mushroom cecid/midge)<br />
Mayetiola avenae (Marchal)<br />
(oat stem midge)<br />
M. destruc<strong>to</strong>r (Say)<br />
(Hessian fly)<br />
Resseliella <strong>the</strong>obaldi (Barnes)<br />
(raspberry cane midge)<br />
* Rhopalomyia chrysan<strong>the</strong>mi (Ahlberg)<br />
(Chrysan<strong>the</strong>mum midge)<br />
Si<strong>to</strong>diplosis mose/lana (Gehin)<br />
(wheat midge)<br />
PHORIDAE<br />
* Megaselia bovista (Gimmerthal)<br />
(mushroom flies)<br />
*M. ha/terata (Wood)<br />
(mushroom flies)<br />
Mushroom<br />
Dit<strong>to</strong>, also cucumber in greenhouses<br />
Dit<strong>to</strong><br />
Cucumber seedlings in greenhouses<br />
Pota<strong>to</strong>; <strong>to</strong>ma<strong>to</strong> and cucumber seedlings in greenhouses<br />
Red clover<br />
Hops<br />
Swede, cabbage, cauliflower, rape, kale, turnip,<br />
radish<br />
Peas (pod & pea)<br />
Pear<br />
Wheat; rye, barley<br />
Rape, cabbage, turnip, radish, mustard, swede,<br />
beet<br />
Red and white clover flowers<br />
Blackcurrant<br />
White clover leaves<br />
Wheat, barley<br />
Mushrooms<br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
Oats<br />
Wheat; barley, rye<br />
Raspberry<br />
Chrysan<strong>the</strong>mum<br />
Rye, oats or barley<br />
Mushroom<br />
Dit<strong>to</strong><br />
12
PHORIDAE (cont.)<br />
*M. nigra (Meigen)<br />
(mushroom flies)<br />
Dit<strong>to</strong><br />
SYRPHIDAE<br />
* Eumerus strigatus (Fallen)<br />
(lesser bulb/narcissus fly)<br />
*E. tubercula/us Rondani<br />
(lesser bulb/narcissus fly)<br />
.Werodon equestris (Fabricius)<br />
(large bulb/narcissus fly)<br />
Bulbs <strong>of</strong> Narcissus. Amaryllis, snowdrops, etc.,<br />
occasionally onion, parsnip, pota<strong>to</strong><br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
TEPHRITIDAE<br />
Euleia ( = Acidia) heracleii (Linnaeus)<br />
(celery fly)<br />
Platyparea poeciloptera (Schrank)<br />
(asparagus fly)<br />
Celery, parsnip<br />
Asparagus (no recent records)<br />
PSILIDAE<br />
Psi/a nigricornis Meigen<br />
(Chrysan<strong>the</strong>mum s<strong>to</strong>ol miner)<br />
* Ps. rosae (Fabricius)<br />
(carrot fly)<br />
Chrysan<strong>the</strong>mum; carrot, lettuce<br />
Carrot, also parsnip, celery, parsley, turnip<br />
OPOMYZIDAE<br />
Geomyza tripunctata Fallen<br />
(grass & cereal fly)<br />
Opomyzajlorum (Fabricius)<br />
(grass & cereal fly)<br />
Grasses; occasionally wheat<br />
Grasses; cereals, especially wheat<br />
EPHYDRIDAE<br />
Hydrellia griseola (Fallen) & spp.<br />
(cereal leaf-miners)<br />
H. nasturtii Collin<br />
(watercress stem miner)<br />
Grasses; cereals (see Deonier, 1971: I 06)<br />
Watercress<br />
DROSOPHILIDAE<br />
Scap<strong>to</strong>myza apica/is Hardy<br />
:\GROMYZIDAE<br />
. .fgromyza nigrella Rondani<br />
(cereal leaf miner)<br />
_.f _ rondensis Strobl<br />
(cereal leaf miner)<br />
Liriomyza bryoniae (Kaltenbach)<br />
(<strong>to</strong>ma<strong>to</strong> leaf miner)<br />
L congest a (Becker)<br />
(pea leaf miner)<br />
L pisivora Hering<br />
(pea leaf miner)<br />
Ophiomyia simplex (Loew)<br />
Phy<strong>to</strong>bia cerasiferae (Kangas)<br />
Phy<strong>to</strong>myza horticola Goureau<br />
P. rujipes Meigen<br />
P. syngenesiae (Hardy)<br />
Broccoli, sprouts, cauliflower, turnip, radish &<br />
watercress; occasionally damages swede, rape,<br />
kale, mustard<br />
Winter wheat<br />
Dit<strong>to</strong><br />
Glasshouse <strong>to</strong>ma<strong>to</strong>, cucumber<br />
Pea and bean<br />
Pea<br />
Asparagus<br />
Plum trees<br />
Pea and bean<br />
Cauliflower, cabbage, broccoli, rape<br />
Chrysan<strong>the</strong>mum<br />
13
CHLOROPIDAE<br />
Ch/orops pumilionis (Bjerkander)<br />
( = taeniopus Meigen)<br />
(gout fly)<br />
Merornyza sa/tatrix (Linnaeus)<br />
(grass fly)<br />
*Oscinellafrit (Linnaeus)<br />
(frit fly)<br />
HYPODERMATIDAE<br />
*Hypoderma bovis (Linnaeus)<br />
(warble fly)<br />
H. diana Brauer<br />
(deer warble fly)<br />
H. lineatum (Villers)<br />
(lesser warble fly)<br />
OESTRIDAE<br />
Oestrus ovis Linnaeus<br />
(sheep nostril fly)<br />
GASTEROPHILIDAE<br />
Gasterophilus haemorrhoidalis (Linnaeus)<br />
(nose bot fly)<br />
G. intestina/is (Degeer)<br />
(horse bot fly)<br />
G. nasalis (Linnaeus)<br />
(throat bot fly)<br />
G. pecorurn (Fabricius)<br />
CALLIPHORIDAE<br />
*Calliphora vicina Robineau-Desvoidy<br />
(bluebottle)<br />
C. vomi<strong>to</strong>ria (Linnaeus)<br />
(bluebottle)<br />
Lucilia caesar (Linnaeus)<br />
(green bottle)<br />
* L. sericata (Meigen)<br />
(sheep maggot fly)<br />
SCATHOPHAGIDAE<br />
Nanna ( = Arnaurosorna) spp.<br />
(timothy flies)<br />
Norellia spinipes (Meigen)<br />
ANTHOMYIIDAE<br />
* Delia antiqua (Meigen)<br />
(onion fly)<br />
*D. radicurn (L.)<br />
( = brassicae) (Wiedemann)<br />
(cabbage root fly)<br />
D. cardui (Meigen)<br />
(carnation fly)<br />
*D. coarctata (Fallen)<br />
(wheat bulb fly)<br />
D. echinata (Seguy)<br />
(spinach stem fly)<br />
D.jloralis (Fallen)<br />
(turnip root fly)<br />
Barley; wheat, rye<br />
Grasses; wheat, barley, rye<br />
Oats; sweetcorn, barley, wheat, rye, maize, ryegrass,<br />
meadow fescue<br />
Cattle, occasionally horses<br />
Deer<br />
Cattle, occasionally horses<br />
Sheep, goats<br />
Horse, donkey<br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
Dit<strong>to</strong><br />
Sheep (secondary)<br />
Dit<strong>to</strong><br />
Sheep (occasional)<br />
Sheep<br />
Timothy grass; winter rye<br />
Narcissus (daffodils)<br />
Onion, leek, shallot, tulip<br />
Cabbage, cauliflower, sprouts, kale, radish,<br />
turnip, swede, mustard, beet, celery & s<strong>to</strong>cks<br />
Carnation<br />
Wheat, rye, barley<br />
Spinach<br />
Turnip, swede<br />
14
ANTHOMYIIDAE (cont.)<br />
D. platura (Meigen)<br />
(bean seed fly)<br />
Pegohy/emyia gnava (Meigen)<br />
(lettuce seed fly)<br />
• Pegomya hyoscami (Panzer)<br />
(beet leaf miner)<br />
P. rubivora (Coquille!!)<br />
(loganberry cane fly)<br />
Phorbia securis (Tiensuu)<br />
(late wheat shoot fly)<br />
P. sepia (Meigen)<br />
(late wheat shoot fly)<br />
Bean, turnip, beet, radish, swede, pota<strong>to</strong>, onion,<br />
pea, cabbage, sprouts, cauliflower, lettuce, rye,<br />
clover, oats, wheat<br />
Lettuce<br />
Mangold, beet, sugarbeet, spinach<br />
Loganberry, raspberry, blackberry<br />
Wheat<br />
Wheat, oats<br />
Advisory leaflets on <strong>the</strong> more important species (marked *above) and some under<br />
crops (e.g. Brassica, mushrooms) are available from <strong>the</strong> Ministry <strong>of</strong> Agriculture,<br />
Fisheries and Food (Publications), Tolcarne Drive, Pinner, Middlesex HAS 2DT.<br />
In addition <strong>to</strong> <strong>the</strong> species whose larvae attack food crops listed above several Cecidomyiidae<br />
are occasional pests <strong>of</strong> grasses, trees and shrubs (especially willows, used for<br />
cricket bats and basket making) and ornamental plants (see Barnes, H. F., 1946-1956);<br />
several o<strong>the</strong>r Agromyzidae also attack garden plants (see Spencer, 1973b ).<br />
Diptera larvae may be found in buildings associated with food production. Several<br />
flies breed in poultry houses and <strong>the</strong> most troublesome is Fannia canicularis Linnaeus<br />
(Fanniidae) though Muscidae (Musca domestica Linnaeus, Muscina stabulans<br />
(Fallen) and Ophyra spp.) also occur. Slaughterhouses are frequented by <strong>the</strong> carrion<br />
flies Lucilia, Calliphora and Sarcophaga. Kiihlhorn (1964) found 330 species <strong>of</strong> flies in<br />
47 families associated with stables. Mushroom houses yield Sciaridae and Phoridae.<br />
\faggots (gentles) bred commercially for use by fishermen include Calliphora and<br />
Pro<strong>to</strong>phormia.<br />
Species specifically associated with excrement are discussed under life-his<strong>to</strong>ries and<br />
ecology.<br />
O<strong>the</strong>r families <strong>of</strong> Diptera are <strong>of</strong> some agricultural (including veterinary) and<br />
economic importance, but not as larvae, e.g. Cera<strong>to</strong>pogonidae, Tabanidae, Muscidae,<br />
Hippoboscidae.<br />
Medical, environmental health and veterinary importance<br />
The larvae <strong>of</strong> species which suck blood or transmit disease as adults can be <strong>of</strong> indirect<br />
medical or veterinary importance, e.g. Cera<strong>to</strong>pogonidae, Culicidae, Simuliidae and<br />
Tabanidae. In <strong>the</strong>se groups (especially abroad) it is <strong>of</strong>ten more effective <strong>to</strong> direct<br />
control measures against <strong>the</strong> larvae (mostly aquatic) ra<strong>the</strong>r than <strong>the</strong> adults. Because <strong>of</strong><br />
this <strong>the</strong> <strong>immature</strong> <strong>stages</strong> have been intensively studied and <strong>the</strong>ir taxonomy and ecology<br />
are better known than in o<strong>the</strong>r families (see Smith, 1973). The larvae <strong>of</strong> o<strong>the</strong>r families<br />
may be more directly involved with man.<br />
Myiasis<br />
Diptera larvae sometimes feed on or in living animals including man and this condition<br />
is called myiasis, a term proposed by Hope (1840) in his early treatise on <strong>the</strong><br />
subject. Zumpt (1965) fully covers <strong>the</strong> subject for <strong>the</strong> Old World, James (1947) for <strong>the</strong><br />
~World and Smith (1973, world) is useful. Only <strong>the</strong> (much rarer) <strong>British</strong> cases<br />
io\·olving man are cited in detail here though all <strong>the</strong> families involved are listed. No<br />
15
doubt many more cases occur than are diagnosed and recorded. Where myiasis is a<br />
normal way <strong>of</strong> life for fly larvae in animals this is described under <strong>the</strong> particular Diptera<br />
family concerned. The types <strong>of</strong> myiasis can be fur<strong>the</strong>r classified as follows.<br />
1. Blood-sucking maggots. E.g. Calliphoridae (Auchmeromyia senegalensis<br />
Macquart( = luteola Fabricius), <strong>the</strong> Congo floor-maggot (Afro-tropical)); Tabanidae<br />
(in paddy fields in Japan); Therevidae (fig. 161, Smith, 1979a, 1986a, <strong>British</strong> cases);<br />
Pro<strong>to</strong>calliphora on birds.<br />
2. Dermal and subdermal myiasis. Larvae penetrate unbroken skin or enter wounds<br />
or form boils or o<strong>the</strong>r lesions, ei<strong>the</strong>r at <strong>the</strong> site <strong>of</strong> entry or elsewhere, e.g. Calliphoridae,<br />
Gasterophilidae, Hypodermatidae, Muscidae and Sarcophagidae.<br />
Oldroyd (1964) reports 'I have known a female Sarcophaga <strong>to</strong> fly <strong>to</strong> a coat, <strong>the</strong> sleeve<br />
<strong>of</strong> which had been soiled with blood from a shot pheasant and immediately lay a batch<br />
<strong>of</strong> young larvae upon it'. Lucilia sericata (Meigen), a common green bottle, is also <strong>the</strong><br />
sheep maggot fly (fig. 809), <strong>the</strong> causal agent <strong>of</strong> sheep strike. Larvae <strong>of</strong> Lucilia and<br />
Calliphora may occur in cases <strong>of</strong> wound myiasis in man and animals. Calliphorid larvae<br />
(Lucilia sericata, L. caesar (Linnaeus) and Phormia regina (Meigen) have been used<br />
<strong>to</strong> clean wounds <strong>of</strong> necrotic tissue on <strong>the</strong> battlefield (Hinman, 1933, Imms, 1939,<br />
Greenberg, 1973) and may even exude an antibiotic (Pavillard & Wright, 1957). Osteomyelitis<br />
and o<strong>the</strong>r kinds <strong>of</strong> non-healing wounds have also been treated in this manner,<br />
e.g. heat burns, X-ray burns, varicose and diabetic ulcers, etc. (see also below under<br />
Musca domestica).<br />
First ins tar larvae <strong>of</strong> Gasterophilus can cause a creeping myiasis in man, usually on<br />
<strong>the</strong> face and but<strong>to</strong>cks (Zumpt, 1965) but I know <strong>of</strong> no <strong>British</strong> cases. Hypoderma<br />
may also be incriminated in this category (Smart, 1939) but is usually involved in<br />
nasopharyngeal myiasis (see below).<br />
Musca domestica larvae may occur in cases <strong>of</strong> human myiasis where urine is<br />
involved: e.g. in neglected cots or napkins <strong>of</strong> infants (Mumford, 1926, Chapman, R. K.,<br />
1944); in <strong>the</strong> feet and footwear <strong>of</strong> incontinent geriatric patients (see also urogenital<br />
myiasis). M . domestica has also been used <strong>to</strong> clean gangrenous wounds as <strong>the</strong> larvae do<br />
not attack healthy tissue (as o<strong>the</strong>r genera will- see above). Healing can commence<br />
48 hours after treatment begins (Imms, 1939).<br />
3. Nasopharyngeal myiasis. This includes invasion <strong>of</strong> <strong>the</strong> nasal fossae, frontal<br />
sinuses, pharyngeal cavities, eyeballs, eye-sockets and (aural myiasis) ears, e.g. Calliphoridae,<br />
Cuterebridae (non-<strong>British</strong>), Gasterophilidae, Hypodermatidae, Myce<strong>to</strong>philidae,<br />
Oestridae, Phoridae, Psychodidae, Sarcophagidae and Scenopinidae.<br />
Lucilia has been involved in a case <strong>of</strong> human aural myiasis in <strong>the</strong> U.K. (Smart, 1936)<br />
and several o<strong>the</strong>r cases are known from Europe and North America. Occasionally first<br />
instar larvae (fig. 693) <strong>of</strong> Hypoderma (mostly lineatum Villers) (warble flies <strong>of</strong> cattle)<br />
have been found in <strong>the</strong> orbit or eyeball <strong>of</strong> man, which <strong>the</strong>y will completely destroy if<br />
<strong>the</strong>y are not extracted quickly. I know <strong>of</strong> only two <strong>British</strong> cases involving H . lineatum,<br />
one in Devon (Style, 1924)and <strong>the</strong> o<strong>the</strong>r in <strong>the</strong> Hope Valley <strong>of</strong> Derbyshire in 1945 when<br />
this locality suffered '<strong>the</strong> worst "gadfly" epidemic for many years' (Smith, A. &<br />
Greaves, 1946) (<strong>the</strong> term 'gadding' <strong>of</strong> cattle is properly used in connection with <strong>the</strong>ir<br />
response <strong>to</strong> <strong>the</strong> attention <strong>of</strong> <strong>the</strong> blood-sucking adult Tabanidae). The larvae <strong>of</strong> Oestrus<br />
ovis Linnaeus (<strong>the</strong> sheep nasal bot fly, fig. 684) may also occur in <strong>the</strong> human eye, but are<br />
far less serious than Hypoderma since <strong>the</strong>y do not survive beyond <strong>the</strong> first instar.<br />
<strong>An</strong>derson (1935) found a dead first instar larva <strong>of</strong> Gasterophilus intestinalis (De Geer)<br />
in <strong>the</strong> eye <strong>of</strong> a patient (in <strong>the</strong> U.S.A.).It is possible that an egg was transferred by wiping<br />
<strong>the</strong> back <strong>of</strong> <strong>the</strong> hand across <strong>the</strong> eye. Zumpt (1965) records (in Africa) seeing G. intestinalis<br />
repeatedly ovipositing on <strong>the</strong> backs <strong>of</strong> human hands when held with <strong>the</strong> palms<br />
over horses' lips.<br />
16
Hurd (1954) records an unique case <strong>of</strong> 13 larvae <strong>of</strong> Boletina (Myce<strong>to</strong>philidae) and<br />
several o<strong>the</strong>r insects being passed alive from <strong>the</strong> left atrum <strong>of</strong> his sinus following<br />
prolonged use <strong>of</strong> <strong>the</strong> aspira<strong>to</strong>r or 'pooter' for insect collecting.<br />
The occurrence <strong>of</strong> Psychodidae in nasopharyngeal myiasis involves <strong>the</strong> non-<strong>British</strong><br />
Clogmia ( = Telma<strong>to</strong>scopus) albipunctatus Willis<strong>to</strong>n (Mohammed & Smith, 1976) but<br />
<strong>British</strong> species occurring in similar habitats (outdoor washtubs, etc.) could be similarly<br />
incriminated (e.g. Psychoda alternata Say, Ps. albipennis Zetterstedt) already recorded<br />
in case <strong>of</strong> intestinal and urogenital myiasis. The accidental inhaling <strong>of</strong> gravid females <strong>of</strong><br />
<strong>the</strong>se small flies through <strong>the</strong> mouth or nostrils could have given rise <strong>to</strong> some <strong>of</strong> <strong>the</strong>se<br />
cases and is <strong>the</strong> probable explanation for <strong>the</strong> unique case <strong>of</strong> lung myiasis produced by<br />
Phoridae reported in a Japanese student and runner (Komori et al. , 1978).<br />
Larvae <strong>of</strong> Lucilia, Fannia, Sarcophaga and Oestrus have been recorded in (rare) cases<br />
<strong>of</strong> aural miasis.<br />
4. Intestinal (enteric) myiasis. This category includes larvae adapted <strong>to</strong> living as<br />
obliga<strong>to</strong>ry parasites in <strong>the</strong> intestines, e.g. Gasterophilidae (horse bot-flies, see under<br />
that family) and those that may be accidentally swallowed in food and be passed in <strong>the</strong><br />
faeces and some which may invade <strong>the</strong> intestine via <strong>the</strong> rectum as facultative parasites,<br />
e.g. <strong>An</strong>isopodidae, Calliphoridae, Drosophilidae, Ephydridae, Fanniidae, Micropezidae,<br />
Muscidae, Phoridae, Piophilidae, Psychodidae, Sarcophagidae, Sepsidae,<br />
Sphaeroceridae, Stratiomyidae, Syrphidae, Therevidae, Tipulidae.<br />
It is improbable that <strong>the</strong> majority <strong>of</strong> ingested fly larvae can survive in <strong>the</strong> digestive<br />
tract when this is in a normal state. However, <strong>the</strong> ingestion <strong>of</strong> larvae <strong>of</strong>ten causes<br />
diarrhoea and <strong>the</strong> larvae are <strong>the</strong>n passed out alive in <strong>the</strong> faeces. Some larvae seem<br />
especially able <strong>to</strong> survive such conditions (e.g. Piophilidae, Phoridae). True rectal<br />
myiasis may occur when flies are attracted <strong>to</strong> excreta and lay eggs on or near <strong>the</strong> anus.<br />
Subsequently larvae may penetrate <strong>the</strong> posterior part <strong>of</strong> <strong>the</strong> rectum and obtain oxygen<br />
by placing <strong>the</strong>ir posterior spiracles in <strong>the</strong> anal region (causing irritation <strong>the</strong>re). Larvae<br />
<strong>of</strong> Eristalis are well adapted <strong>to</strong> this mode <strong>of</strong> life and <strong>the</strong>re are many records <strong>of</strong> <strong>the</strong>ir<br />
involvement in myiasis including Britain (Mumford, 1926; Cookson & Oldroyd, 1937).<br />
Fannia canicularis (Linnaeus) and F. scalaris (Fabricius) have also been involved in<br />
rectal myiasis. The larvae <strong>of</strong> Sarcophaga may cause true rectal myiasis and Zumpt<br />
( 1965) records larvae <strong>of</strong> S. cruentata Meigen ( = haemorrhoidalis (Fallen)) seen leaving<br />
and re-entering <strong>the</strong> anus <strong>of</strong> a child (in Sydney).<br />
5. Urogenital myiasis. There are no obliga<strong>to</strong>ry dipterous parasites in <strong>the</strong> urogenital<br />
system; all are casual invaders (facultative parasites), e.g. <strong>An</strong>isopodidae, Calliphoridae,<br />
Ephydridae, Fanniidae, Muscidae, Phoridae, Psychodidae, Sarcophagidae,<br />
Syrphidae, Scenopinidae.<br />
Sylvicola ( = <strong>An</strong>isopus) fenestra/is (Linnaeus) larvae have been recorded from <strong>the</strong><br />
urinary tract <strong>of</strong> female (Smith & Taylor, 1966) and male (Morris, R. F., 1968) patients.<br />
Thompson J. H. et al. (1970) record a larva <strong>of</strong> Scenopinus from <strong>the</strong> urine <strong>of</strong> a woman in<br />
<strong>the</strong> U.S.A. James (1947) says <strong>of</strong> Teichomyza fusca Macquart (Ephydridae) that<br />
·numerous cases <strong>of</strong> urinary myiasis have been recorded'; some <strong>of</strong> <strong>the</strong>se are European<br />
but I know <strong>of</strong> no <strong>British</strong> cases (see Fossil section, archaeological sites). Musca domestica<br />
has been involved in cases <strong>of</strong> urogenital myiasis and a fascinating account is given<br />
by Leon (1921) in which a male student in Romania actually ejaculated maggots (also<br />
summarized in Zumpt, 1965: 34).<br />
With <strong>the</strong>ir predilection for urine, it is not surprising that Fannia canicularis and<br />
F. scalaris have frequently been involved in this type <strong>of</strong> myiasis in humans (Haddow<br />
& Lumsden, 1935). Larvae <strong>of</strong> Psychoda albipennis, Ps. alternata, Megaselia sp.,<br />
Calliphora vicina and Sarcophaga species have also been reported as causing urogenital<br />
myiasis.<br />
Eristalis has been recorded from <strong>the</strong> vagina <strong>of</strong> a cow (and tinned pears!).<br />
17
Forensic or medico-legal importance<br />
After death animal and human corpses are invaded by a succession <strong>of</strong> insects and<br />
o<strong>the</strong>r invertebrates associated with <strong>the</strong> various <strong>stages</strong> <strong>of</strong> decomposition. Not surprisingly,<br />
many <strong>of</strong> <strong>the</strong> Diptera involved are <strong>of</strong> <strong>the</strong> same families, and frequently <strong>the</strong> same<br />
species, as those involved in myiasis. The most important decomposers are blowfly<br />
(Calliphoridae) larvae <strong>of</strong> <strong>the</strong> genera Calliphora and Lucilia.<br />
Identification <strong>of</strong> <strong>the</strong> species involved, combined with a knowledge <strong>of</strong> <strong>the</strong>ir rates <strong>of</strong><br />
development and ecology, enables an estimate <strong>of</strong> time <strong>of</strong> death <strong>to</strong> be made. It may also<br />
be possible <strong>to</strong> establish if a body has been moved or concealed for part <strong>of</strong> <strong>the</strong> time. The<br />
faunal succession on carrion is dealt with in <strong>the</strong> section on life-his<strong>to</strong>ries and ecology and<br />
<strong>the</strong> forensic aspects are fully covered elsewhere (Smith, 1986b ).<br />
<strong>An</strong>o<strong>the</strong>r important medico-legal involvement with larvae is <strong>the</strong>ir presence on foodstuffs,<br />
drugs, etc. in domestic or commercial premises. Frequently en<strong>to</strong>mologists are<br />
asked, usually by environmental health <strong>of</strong>ficers, not only <strong>to</strong> identify such larvae but <strong>to</strong><br />
give additional information on <strong>the</strong> age <strong>of</strong> <strong>the</strong> larva, its country <strong>of</strong> origin and whe<strong>the</strong>r it<br />
could have survived refrigeration. This enables <strong>the</strong> possible source and time <strong>of</strong> infestation<br />
<strong>to</strong> be estimated and blame <strong>to</strong> be apportioned in cases involving prosecution.<br />
Bus vine ( 1980) is particularly valuable for this type <strong>of</strong> enquiry and Oldroyd (in Smith,<br />
1973) is useful. Such enquiries may involve larvae <strong>of</strong> non-<strong>British</strong> species and <strong>the</strong> larvae<br />
<strong>of</strong> <strong>the</strong> commonest <strong>of</strong> <strong>the</strong>se are included in <strong>the</strong> present Handbook, e.g. Ceratitis capitata<br />
(Wiedemann) (Mediterranean fruit fly, in oranges, lemons, mangos, etc.), Rhago/etis<br />
cerasi (Linnaeus) (cherry fruit-fly in cherries and cherry yoghurt) (both Tephritidae);<br />
Hermetia il/ucens (Linnaeus) (Stratiomyidae, in tinned <strong>to</strong>ma<strong>to</strong>es, black peppers, root<br />
ginger, mango slices in brine, etc.) (Smith & Chainey, 1989).<br />
Some common types <strong>of</strong> 'economic' enquiries involving native <strong>British</strong> species (plus<br />
Hermetia illucens) are listed below with <strong>the</strong> fly larvae involved in order <strong>of</strong> frequency <strong>of</strong><br />
occurrence in cases I have dealt with over a period <strong>of</strong> more than 30 years.<br />
A. Larvae in foods, drinks, etc.<br />
I. Meat and meat products, poultry, fish, etc.: Calliphora, Lucilia, Hermetia illucens, Sarcophaga,<br />
Hypoderma (in venison).<br />
2. Cheese: Calliphora (also butter), Piophi/a, Fannia (sometimes in wrappings <strong>of</strong> o<strong>the</strong>r animal<br />
protein foods, e.g. butter), Sarcophaga.<br />
3. Eggs, broken and bad: Muscina (Muscidae), Megaselia (Phoridae).<br />
4. Milk bottles (empty or full); Spiniphora bergenstammi (Mik) (Phoridae), Drosophila (especially<br />
D. funebris (Fabricius) and D. busckii Coquillett) (Drosophilidae), Calliphora, Muscina.<br />
Psychodidae (Psychoda surcoufi Tonnoir), Syrphidae (Syrphini, Eristalis), Trichocera<br />
(Trichoceridae).<br />
5. Vegetables & vegetable products: Phoridae, Sciaridae, Hermetia illucens, Syrphidae,<br />
Lonchopteridae, Chironomidae (Metriocnemus in salads including watercress).<br />
6. Fruit & fruit products: Eristalis, Tephritidae, Drosophilidae, Hermetia illucens, Muscina<br />
stabu/ans (Fallen).<br />
7. Alcoholic & fruit drinks, vinegar, etc.: Drosophilidae larvae may still be recognisable in dehydrated<br />
food after reconstitution (see Sciaridae).<br />
B. Larvae in domestic premises<br />
I. Larvae emerging from soil, cracks in concrete, etc. (especially after heavy rain), in numbers on<br />
concrete surfaces or indoors: Tipu/a, Lucilia.<br />
2. Indoors (including upstairs rooms): <strong>Vol</strong>ucella (from wasps' nests in l<strong>of</strong>t), puparia <strong>of</strong> Crataerina<br />
pallida (from swift and house-martin nests), Calliphora (from nearby corpse (bird, mouse, etc.<br />
in l<strong>of</strong>t, chimney, under floorboards or carpets)), Scenopinus (in or under carpets), Syrphus<br />
(from leaves <strong>of</strong> cut flowers, salads, etc.), Sciaridae in neglected houseplant pots (and Oldroyd<br />
(1964) records a bizarre case where sciarid larvae infested <strong>the</strong> mouldy peaty material used as a<br />
fire-resisting lining in an <strong>of</strong>fice safe).<br />
18
3. In water supply: Chironomidae, Psychodidae, Eristalis (Syrphidae, if water is contaminated).<br />
4. Faulty or blocked WCs: Lep<strong>to</strong>cera caenosa (Rondani), Fannia, Musca domestica, Eristalis,<br />
Psychodidae.<br />
5. In refuse in dustbins: Calliphora, Luci/ia.<br />
In food products where meat and vegetables (and sauces, pickles, etc.) are mixed,<br />
some transference may occur <strong>to</strong> complicate <strong>the</strong> picture on <strong>the</strong> plate or in <strong>the</strong> container<br />
(e.g. one case where jets <strong>of</strong> Drosophila larvae were ejected from a plastic <strong>to</strong>ma<strong>to</strong><br />
ketchup dispenser!).<br />
Larvae <strong>of</strong> species specific <strong>to</strong> particular vegetable crops in <strong>the</strong> field may occasionally<br />
be transported alive indoors and crawl or drop <strong>of</strong>f <strong>the</strong> foodplant (e.g. De/ia radicum L.<br />
( = brassicae), <strong>the</strong> cabbage root-fly, or aphidophagous Syrphidae). These and o<strong>the</strong>rs<br />
are included under species <strong>of</strong> agricultural importance.<br />
Larvae <strong>of</strong> Calliphora on leaving <strong>the</strong>ir pabulum <strong>to</strong> search for suitable pupation sites<br />
may occur in a variety <strong>of</strong> indoor situations, e.g. bedrooms (from bird corpse in 'sealed'<br />
fireplace), in freshly delivered rolls <strong>of</strong> carpet!<br />
Adults <strong>of</strong> several families frequently occur in numbers indoors and a consideration<br />
<strong>of</strong> <strong>the</strong>ir life-his<strong>to</strong>ries may provide clues <strong>to</strong> <strong>the</strong>ir source <strong>of</strong> origin.<br />
Preda<strong>to</strong>rs, parasites and pathogens<br />
"The difference between ... a carnivore and a parasite is simply <strong>the</strong> difference<br />
between living upon capital and income."<br />
Charles El <strong>to</strong>n, 1927<br />
Man certainly plays a large part in controlling <strong>the</strong> numbers <strong>of</strong> flies by attacking <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> ei<strong>the</strong>r deliberately in controlling economic pests, as food (not intentionally<br />
in U.K.), or accidentally by his modification <strong>of</strong> <strong>the</strong> environment. However, he<br />
also provides many artificial habitats which favour <strong>the</strong> development <strong>of</strong> synanthropic<br />
Diptera. More specific enemies may be classified as preda<strong>to</strong>rs or parasites.<br />
Preda<strong>to</strong>rs<br />
Diptera larvae in <strong>the</strong> soil probably form a regular part <strong>of</strong> <strong>the</strong> diets <strong>of</strong> mammals with a<br />
5Ubterranean life style: moles, for example, take larvae <strong>of</strong> Tipulidae, <strong>An</strong>isopodidae,<br />
Bibionidae, Sciaridae, Stratiomyidae, Therevidae, Rhagionidae, Tabanidae, Empididae<br />
and <strong>An</strong>thomyiidae (Godfrey & Crowcr<strong>of</strong>t, 1960). Birds will take larvae near <strong>the</strong><br />
surface, and lea<strong>the</strong>rjackets, for example, are eaten by rooks, starlings, plover, gulls<br />
and pheasants (Smith, K. M., 1931 ). Carnivorous soil-dwelling invertebrates such as<br />
centipedes also take fly larvae as food. Some soil-dwelling beetles such as Carabidae<br />
and Staphylinidae are carnivorous as adults and larvae and feed on Diptera larvae. The<br />
saaphylinid Aloconota gregaria Erichson may be a significant biological control agent<br />
against <strong>the</strong> wheat bulb fly Delia coarctata (Fallen). Predaceous Diptera larvae in<br />
tbe soil may prey on o<strong>the</strong>r Diptera larvae including each o<strong>the</strong>r, e.g. Empididae,<br />
Dolichopodidae (Hobby & Smith, 1961 b). Many beetles, for example Staphylinidae,<br />
Silpbidae and Cleridae, prey on fly eggs and larvae in carrion, fungi, etc. Wasps take<br />
Diptera larvae in exposed situations on carrion, etc.<br />
Aquatic Diptera larvae are subject <strong>to</strong> heavy predation from fish and some water<br />
birds (e.g. dippers, ducks) as well as from <strong>the</strong> orders <strong>of</strong> insects with aquatic carnivorous<br />
lanae, such as Odonata, Plecoptera, Hemiptera and Coleoptera. Larvae <strong>of</strong> <strong>the</strong><br />
~nomid genera Cardiocladius and Eukiefferiella will attack Simulium. Larvae <strong>of</strong><br />
_.falfttlrocnema (C/inoceroides) glaucescens (Loew) (Scathophagidae) feed on egg<br />
19
masses <strong>of</strong> Dixa (Dixidae) and larvae <strong>of</strong> Phaonia exoleta (Meigen) (Muscidae) feed on<br />
mosqui<strong>to</strong> (Culicidae) larvae.<br />
Some may also be attacked by adult Diptera, e.g. chironomid larvae are attacked by<br />
adult Dolichopodidae (Hydrophorus, Campsicnemus, Poecilobothrus and Dolichopus)<br />
(Smith & Empson, 1955). Simuliid larvae have been reported <strong>to</strong> be attacked by adult<br />
Empididae (Ciinocerinae and Hemerodromiinae) and Dolichopodidae in Canada<br />
(Davies, D . M., 1981).<br />
On plants, predaceous larvae <strong>of</strong> Les<strong>to</strong>diplosis are commonly found feeding on larvae<br />
<strong>of</strong> o<strong>the</strong>r Cecidomyiidae (Harris, 1966).<br />
In <strong>the</strong> U.S.A. water mites have been recorded as preda<strong>to</strong>rs on <strong>the</strong> eggs and larvae <strong>of</strong><br />
mosqui<strong>to</strong>es (Mullen, 1975).<br />
Parasites<br />
Parasites <strong>of</strong> Diptera should always be carefully preserved with all <strong>the</strong> relevant<br />
puparial fragments, etc. and data, as much remains <strong>to</strong> be discovered and many rea rings<br />
in <strong>the</strong> literature are in need <strong>of</strong> careful confirmation.<br />
The Hymenoptera provide <strong>the</strong> most important parasites <strong>of</strong> <strong>immature</strong> Diptera. A<br />
general survey is given by Shaw & Askew (in Stubbs & Chandler, 1978) and Smith<br />
(1974b). The principal parasites are as follows: Ichneumonidae (Phygadeuon (fig.<br />
1321), Mesoleptus and related genera on fly puparia, Oxy<strong>to</strong>rinae on Myce<strong>to</strong>philidae,<br />
Orthocentrinae on Diptera near <strong>the</strong> ground, Diplazontinae on aphidophagous<br />
Syrphidae): Braconidae (Opiinae on plant feeding Schizophora, Alysiinae (figs 1319,<br />
1320) on Cyclorrhapha); Eulophidae; Pteromalidae; some Ceraphron<strong>to</strong>idea; Proc<strong>to</strong>trupoidea<br />
(Piatygastridae on Cecidomyiidae); Diapriidae (Diapriinae on cyclorrhaphous<br />
puparia, Belytinae on Myce<strong>to</strong>philidae), Scelionidae, Trichogrammatidae,<br />
Mymaridae (all <strong>of</strong> <strong>the</strong> last three families are egg parasites on some Cecidomyiidae,<br />
Tabanidae, Rhagionidae and Syrphidae); Cynipoidea (Eucoilidae and Figitidae on<br />
Cyclorrhapha).<br />
In <strong>the</strong> Coleoptera some larvae <strong>of</strong> Staphylinidae, e.g. Aleochara, are parasites <strong>of</strong> fly<br />
puparia, mostly calypterates and especially <strong>An</strong>thomyiidae (see Welch, 1965). The<br />
species <strong>of</strong> fly parasitised probably varies with <strong>the</strong> habitat, e.g. Aleochara algarum<br />
Fauvel is parasitic on seaweed flies such as Coelopa and Orygma, while A. cur tu/a<br />
(Goeze) is a common parasite in <strong>the</strong> puparia <strong>of</strong> blowflies (figs 1322- 1326).<br />
Parasitism <strong>of</strong> Diptera by Diptera is not common and <strong>the</strong> few <strong>British</strong> records (see<br />
under family for details) may be summarised as follows:<br />
Tachinidae: Siphona on lea<strong>the</strong>rjacket larvae <strong>of</strong> Tipulidae and a Pegomya sp.;<br />
Admontia (Trichopareia) specialises in Tipulidae in wood, e.g. Ctenophora, Tipula<br />
irrorata Macquart and T.jlavolineata Meigen.<br />
Phoridae: Borophaga incrassata Meigen parasitizes larvae <strong>of</strong> Bibio marci L.; M egaselia<br />
paludosa Wood attacks lea<strong>the</strong>rjackets; M .jlavicoxa Zetterstedt parasitises larvae <strong>of</strong><br />
Bradysia bicolor Meigen (Sciaridae); and M. obscuripennis Wood is a parasite <strong>of</strong> larvae<br />
<strong>of</strong> Trichosia (Sciaridae).<br />
In <strong>the</strong> U .S.A. Macronychia (Sarcophagidae) has been reared from adult Tabanidae<br />
and some Tachinidae have been reared from larval and adult Tabanidae (Thompson,<br />
P. H ., 1978). In Australia a tachinid has been reared from an adult A<strong>the</strong>rigona (Muscidae)<br />
(Smith, 1974c, Ferrar, 1977).<br />
Parasitic fungi probably also take a steady <strong>to</strong>ll <strong>of</strong>larvae, as <strong>the</strong>y do <strong>of</strong> adults, though<br />
few are recorded specifically from Diptera larvae in <strong>the</strong> literature (Lea<strong>the</strong>rdale, 1970).<br />
I have found <strong>the</strong> larvae <strong>of</strong> Lonchaea palposa Zetterstedt infested with <strong>the</strong> 'green<br />
muscardine' fungus Metarrhizium anisopliae (Metsch) (Smith, 1957b). D'Arcy-Burt<br />
( 1987) reports <strong>the</strong> hyphomycete Tolypocaladium cylindrosporum Gams from <strong>the</strong> larvae<br />
<strong>of</strong> Bibionidae. Abroad, but not yet in Britain, species <strong>of</strong> Coelomomyces fungi have been<br />
found in <strong>the</strong> larvae <strong>of</strong> mosqui<strong>to</strong>es.<br />
20
Various micro-organisms may infest dipterous larvae, e.g. viruses, bacteria, Pro<strong>to</strong>zoa,<br />
nema<strong>to</strong>des (fig. 1318), ces<strong>to</strong>des and trema<strong>to</strong>des. A general survey is given by<br />
Carter (in Stubbs & Chandler, 1978).<br />
The micro-fauna and flora <strong>of</strong> dipterous larvae is a subject ripe for careful research.<br />
Fossil larvae and pupae<br />
Diptera larvae are sometimes, though rarely, preserved as fossils and, if carefully<br />
examined under <strong>the</strong> microscope in an oblique light, may be identified <strong>to</strong> families. Often<br />
<strong>the</strong>se fossils are ra<strong>the</strong>r faint but details can be enhanced by moistening <strong>the</strong> surface with<br />
a thin film <strong>of</strong> water or 80% alcohol for short periods. However, great care should be<br />
taken that <strong>the</strong> rock containing <strong>the</strong> impression <strong>of</strong> <strong>the</strong> fossil is not soluble (e.g. if gypsum<br />
is present) by testing areas clear <strong>of</strong> <strong>the</strong> fossil. A useful recent reference is Whalley &<br />
Jarzembowski (1985). For fossil Diptera in general Rohdendorf (1974) and Hennig<br />
( 1981) should be consulted.<br />
Many insects (<strong>of</strong>ten fragmented), especially Coleoptera, are found by archaeologists<br />
during <strong>the</strong>ir excavations. The insect fauna on such sites is usually in <strong>the</strong> region <strong>of</strong><br />
prehis<strong>to</strong>ric refuse dumps or cess pits and <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Diptera are frequently<br />
represented as follows: Tipulidae (especially larval head capsules), Chironomidae<br />
(larval head capsules), Drosophilidae (puparia), Sepsidae (puparia), Ephydridae<br />
( Teichomyza and Scatella puparia), Sphaeroceridae (puparia), Scathophagidae<br />
(puparia), Muscidae (Musca, Muscina, Neomyia ( = Or<strong>the</strong>llia), Eudasyphora<br />
( = Dasyphora), puparia or parts <strong>of</strong>).<br />
Puparia are <strong>of</strong>ten darkened and may be covered in debris. They should be soaked in<br />
cold I 0% KOH, cleared <strong>of</strong> debris, <strong>the</strong>n teased apart so that <strong>the</strong> mouthparts can be<br />
extracted from <strong>the</strong> puparial cap (see also collecting, preservation, etc.). Some useful<br />
references for identification are Phipps (1983), Skidmore (1985) and Smith (1986b).<br />
Collecting, rearing, preservation and examination<br />
Dipterists collecting adults usually use nets, and larvae are rarely seen. In fact Diptera<br />
larvae are more likely <strong>to</strong> be encountered by specialists in o<strong>the</strong>r groups <strong>of</strong> insects<br />
using entirely different collecting methods. It is <strong>the</strong>refore well worth while educating<br />
colleagues <strong>to</strong> collect alive any Diptera larvae <strong>the</strong>y may find. This casual method <strong>of</strong><br />
collection should not be spurned as it <strong>of</strong>ten yields Diptera larvae <strong>of</strong> considerable<br />
interest <strong>of</strong> which <strong>the</strong> adults may be only rarely seen.<br />
The systematic collecting <strong>of</strong> larvae does present certain problems as <strong>the</strong> extent <strong>of</strong><br />
possible habitats is so large that some form <strong>of</strong> sampling is necessary. The types <strong>of</strong><br />
habitat occupied by fly larvae are outlined under <strong>the</strong> section on life-his<strong>to</strong>ries and<br />
ecology and <strong>the</strong> collecting methods employed must be dictated by <strong>the</strong> microhabitat<br />
favoured and <strong>the</strong> known, or suspected, behavioural characteristics <strong>of</strong> <strong>the</strong> groups <strong>to</strong> be<br />
collected.<br />
Having collected larvae in a given habitat it is important <strong>to</strong> ascertain where and on<br />
what <strong>the</strong>y were probably feeding. Some larvae occur in strictly defined habitats but<br />
o<strong>the</strong>rs such as certain soil-dwelling larvae may also be found in leaf-litter, wood debris<br />
or dung. Among aquatic larvae some normally associated with running water can be<br />
found in still water and some terrestrial larvae may be found in aquatic moss. The<br />
larvae may move from one microhabitat <strong>to</strong> ano<strong>the</strong>r during <strong>the</strong>ir life-his<strong>to</strong>ry. For<br />
example, semi-aquatic larvae may move <strong>to</strong> drier habitats for <strong>the</strong> purpose <strong>of</strong> pupation,<br />
and carrion-feeding calliphorid maggots migrate from carrion for a considerable distance<br />
prior <strong>to</strong> pupation. All possibilities <strong>of</strong> this sort should be considered when examining<br />
samples and where possible <strong>the</strong> larvae should be grouped in<strong>to</strong> similar types and<br />
21
sizes and some <strong>of</strong> each kept alive for possible rearing <strong>to</strong> adult in order <strong>to</strong> establish<br />
identity. Pupae or puparia should be kept alive.<br />
Glass or plastic tubes or tins are suitable containers for keeping terrestrial larvae<br />
alive, <strong>to</strong>ge<strong>the</strong>r with some <strong>of</strong> <strong>the</strong> pabulum in which <strong>the</strong>y were found. Aquatic larvae<br />
should be placed in suitable jars or bottles. Larvae suspected <strong>of</strong> being carnivorous<br />
should be kept singly.<br />
Useful equipment for collecting terrestrial larvae includes a small trowel for soil<br />
sampling, a strong knife for splitting bark, wood, etc., a sieve or nest <strong>of</strong> sieves <strong>of</strong> graded<br />
mesh size, forceps, a fine paint brush (for tiny delicate larvae), a white sheet or dish for<br />
sorting, a notebook, adhesive labels and pencil. A pooter or insect aspira<strong>to</strong>r could be<br />
useful in some dry situations but must be <strong>of</strong> <strong>the</strong> 'blow' type <strong>to</strong> avoid health hazards (see<br />
section on Medical Importance).<br />
For aquatic species a strong metal-framed pond net is essential; a wire hook for<br />
securing weed is useful and a strong stick for turning s<strong>to</strong>nes and a large white tray for<br />
sorting.<br />
A useful way <strong>of</strong> sorting soil indoors is by floatation in which <strong>the</strong> sample is immersed<br />
and broken up in a solution <strong>of</strong> magnesium sulphate (Epsom salts) at a strength <strong>of</strong> about<br />
one pound (0.488 kilo) <strong>to</strong> one gallon (4.55litres) <strong>of</strong> water. The larvae float on <strong>the</strong><br />
surface (puparia <strong>of</strong>ten around <strong>the</strong> edges) and should be picked <strong>of</strong>f quickly as this<br />
solution can anaes<strong>the</strong>tise aquatic animals. <strong>An</strong> alternative solution can be made using<br />
some common salt in water. The floatation methods only work for free-living larvae<br />
and are no substitute for careful hand searching which may also yield larvae hidden in<br />
cocoons and silken tubes.<br />
Soil-dwelling species may be sampled by using a solution <strong>of</strong> orthodichlorobenzene<br />
(Jeyes' Fluid). About half-an-ounce (14 g) <strong>of</strong> <strong>the</strong> liquid <strong>to</strong> one gallon <strong>of</strong> water is poured<br />
on cleared ground at <strong>the</strong> rate ·<strong>of</strong> 2 gallons (9.1 litres) <strong>of</strong> <strong>the</strong> mixture per square yard<br />
(0.83 sq. m). <strong>An</strong>y larvae should surface before half an hour or so (see Brindle & Smith,<br />
in Stubbs & Chandler, 1978).<br />
Fur<strong>the</strong>r information on collecting and equipment is given in Oldroyd (1970a),<br />
Brindle & Smith (in Stubbs & Chandler, 1978), Cogan & Smith (1974) and Southwood<br />
(1978).<br />
Rearing is only <strong>of</strong> value if <strong>the</strong> larvae or puparia are segregated in<strong>to</strong> suitable individual<br />
containers so that <strong>the</strong> <strong>immature</strong> <strong>stages</strong> can be linked <strong>to</strong> <strong>the</strong> subsequent adult and<br />
identification confirmed. As far as possible <strong>the</strong> conditions in which <strong>the</strong> larvae were found<br />
shou"td be re-created in miniature, especially humidity. Humidity or moisture gradients<br />
can be achieved by tilting containers containing moss, soil and a little water so that subaquatic<br />
larvae can choose <strong>the</strong> level that best suits <strong>the</strong>m. A range <strong>of</strong> plant (or animal) food<br />
collected in <strong>the</strong> larval habitat should be provided as possible food for <strong>the</strong> larvae, care<br />
being taken not <strong>to</strong> introduce anything likely <strong>to</strong> prey upon <strong>the</strong> larva. Close observation<br />
and care is necessary until it is clear that <strong>the</strong> conditions provided are satisfac<strong>to</strong>ry.<br />
Containers can vary from tins, gauze <strong>to</strong>pped tubes or jars, petri dishes, etc., whichever<br />
seems best <strong>to</strong> fit <strong>the</strong> particular moisture requirement without encouraging mould.<br />
Rearing larvae <strong>of</strong> unknown habits and identity is not easy but <strong>the</strong> interest and<br />
satisfaction <strong>of</strong> possibly establishing a new life-his<strong>to</strong>ry and describing a hi<strong>the</strong>r<strong>to</strong><br />
unknown larva is sufficient reward for making <strong>the</strong> attempt. Wong (1972) reviews<br />
rearing techniques.<br />
Larvae selected for preservation are best killed in hot (60-70oC) water if this is<br />
available. This also expands <strong>the</strong> larvae which can <strong>the</strong>n be placed and s<strong>to</strong>red in 80%<br />
alcohol with a little glycerine added <strong>to</strong> guard against drying out through evaporation.<br />
A better medium for preservation is Berlese preservative which is prepared in a similar<br />
way <strong>to</strong> Berlese mountant (see below), but without gum arabic. This is a clear fluid that<br />
does not set and specimens may be kept in it indefinitely and mounted as required. If it<br />
thickens because <strong>of</strong> evaporation it can easily be thinned with small amounts <strong>of</strong> distilled<br />
water.<br />
22
Specimens preserved in alcohol may need <strong>to</strong> be s<strong>of</strong>tened and cleared <strong>to</strong> facilitate<br />
microscopical examination or <strong>the</strong> preparation <strong>of</strong> permanent mounts on microscope<br />
slides. The larvae should be placed in <strong>10</strong>% caustic potash and left overnight or <strong>the</strong><br />
process can be accelerated by placing <strong>the</strong> specimen in caustic potash in a small tube<br />
immersed in boiling water. Very small, delicate larvae should be watched carefully<br />
during this process as <strong>the</strong>y may clear <strong>to</strong>o quickly. The specimens should <strong>the</strong>n be<br />
transferred <strong>to</strong> glacial acetic acid for 5- <strong>10</strong> minutes and finally placed in a drop <strong>of</strong>Berlese<br />
preservative in a watch glass or on a cavity slide for examination or dissection under a<br />
low power stereoscopic microscope. It is better <strong>to</strong> attempt identification before making<br />
a permanent mount as it is <strong>of</strong>ten essential <strong>to</strong> manoeuvre <strong>the</strong> specimen in<strong>to</strong> different<br />
positions. As one proceeds with identification it soon becomes clear from <strong>the</strong> key<br />
characters if dissection will be necessary and if a permanent mount <strong>of</strong> <strong>the</strong> whole or a<br />
part <strong>of</strong> <strong>the</strong> specimen needs <strong>to</strong> be made for more detailed study under a compound<br />
microscope.<br />
Permanent slide mounts can be made by transferring <strong>the</strong> specimen from glacial acetic<br />
acid in<strong>to</strong> clove oil and <strong>the</strong>n in<strong>to</strong> Canada balsam. However <strong>the</strong> refractive index <strong>of</strong><br />
Canada balsam is such that fine structures may be difficult <strong>to</strong> see and a far superior<br />
medium for temporary slides is Swan's Berlese Mountant which is made as follows<br />
(formula after Dr K. M. Harris):<br />
Gum arabic . .<br />
Chloral hydrate .<br />
Glacial acetic acid<br />
50% wjw glucose syrup<br />
Distilled water . . .<br />
12g<br />
20g<br />
5ml<br />
5ml<br />
30-40ml<br />
Measure ingredients in<strong>to</strong> a beaker or wide-mou<strong>the</strong>d bottle (<strong>the</strong> glucose syrup is<br />
made by dissolving a given weight <strong>of</strong> glucose in an equal weight <strong>of</strong> distilled water).<br />
Place in gentle heat (slide-oven at 35°C) for 24 hours or more, stirring occasionally<br />
until ingredients dissolve. Leave for a few days until debris from <strong>the</strong> gum arabic<br />
settles and <strong>the</strong>n decant clear fluid in<strong>to</strong> a clean screw-<strong>to</strong>p bottle. If necessary,<br />
evaporate water by leaving <strong>the</strong> open container in a slide-oven for a few days, until<br />
drops <strong>of</strong> mountant retain <strong>the</strong>ir shape on a slide ra<strong>the</strong>r than spreading over <strong>the</strong><br />
surface.<br />
It is usually necessary <strong>to</strong> examine <strong>the</strong> slides from time <strong>to</strong> time, <strong>to</strong>pping up with<br />
mountant <strong>to</strong> compensate for evaporation. After <strong>the</strong> slides have dried for a few<br />
months (minimum <strong>of</strong> 3 weeks at 35°C) <strong>the</strong>y should be ringed <strong>to</strong> prevent fur<strong>the</strong>r<br />
evaporation. Unfortunately most ringing reagents react with <strong>the</strong> mountant and<br />
crystallization still occurs so that permanent slides should be made with Euparal,<br />
mounting direct from 95 % alcohol or propanol.<br />
If only empty puparia are available <strong>the</strong>y can be treated in <strong>the</strong> same way as larvae,<br />
but it should be remembered that puparial caps may become detached and <strong>the</strong>se are<br />
important since <strong>the</strong>y contain <strong>the</strong> mouthparts <strong>of</strong> <strong>the</strong> last instar larva which can be<br />
used in identification. The spiracles <strong>of</strong> <strong>the</strong> final instar larvae are also present on <strong>the</strong><br />
puparium.<br />
A useful ruse for <strong>the</strong> rapid examination <strong>of</strong> <strong>the</strong> posterior spiracles <strong>of</strong> dead large firm<br />
larvae or puparia (alive or dead) is <strong>to</strong> plunge <strong>the</strong>m head down in some I /4" depth <strong>of</strong> fine<br />
sand in a watch glass which holds <strong>the</strong>m in position under <strong>the</strong> microscope.<br />
Users <strong>of</strong> this Handbook are hardly likely <strong>to</strong> need <strong>the</strong> use <strong>of</strong> <strong>the</strong> scanning electron<br />
microscope. Never<strong>the</strong>less, for <strong>the</strong> clarification <strong>of</strong> minute structures at <strong>the</strong> limit <strong>of</strong> <strong>the</strong><br />
light microscope <strong>the</strong> instrument opens up new dimensions in larval study. The SEM<br />
has transformed our understanding <strong>of</strong> <strong>the</strong> functional morphology <strong>of</strong> egg and larval<br />
structures (e.g. Hin<strong>to</strong>n, 1981). The techniques <strong>of</strong> preparation <strong>of</strong> larval material for<br />
SEM examination are described by Grodowitz et al. (1982).<br />
23
Biology and morphology <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong><br />
Diptera are holometabolous insects with a complete metamorphosis <strong>of</strong> four <strong>stages</strong>:<br />
egg, larva, pupa and adult.<br />
During growth <strong>the</strong> larvae <strong>of</strong> most Diptera pass through a series <strong>of</strong> moults, <strong>the</strong> period<br />
between <strong>the</strong> moults being called ins tars. Moults that take place soon after hatching and<br />
just prior <strong>to</strong> pupation are easily overlooked but those that take place during <strong>the</strong> active<br />
feeding period are easily observed.<br />
Nema<strong>to</strong>cera usually have <strong>the</strong> largest number <strong>of</strong> moults, e.g. four instars in Culicidae<br />
and six or seven in Simuliidae. Among Brachycera, some Tabanidae have up <strong>to</strong> nine<br />
instars and may take up <strong>to</strong> two years <strong>to</strong> complete <strong>the</strong>ir development. The maggots <strong>of</strong><br />
Cyclorrhapha usually have three instars and most feeding and growing is done in <strong>the</strong><br />
third instar.<br />
Some larvae <strong>of</strong> Cecidomyiidae reproduce by paedogenesis, i.e. reproduction by an<br />
animal that becomes sexually mature before reaching <strong>the</strong> adult form . In this method <strong>of</strong><br />
reproduction a few (up <strong>to</strong> about 35) large eggs are produced which can be seen through<br />
<strong>the</strong> larval cuticle. These become larvae and feed on <strong>the</strong>ir parent larva from inside (fig.<br />
118). This condition can continue for several generations without <strong>the</strong> appearance <strong>of</strong> an<br />
adult. These midges (e.g. Mias<strong>to</strong>r) thus reproduce like aphids and remain in <strong>the</strong>ir<br />
birthplace producing a great mass <strong>of</strong>larvae which may be found under <strong>the</strong> bark <strong>of</strong>logs.<br />
Eventually <strong>the</strong> daughter larvae will cease <strong>to</strong> produce eggs and pupate normally. Both<br />
sexes are produced in this way though <strong>of</strong> course all <strong>of</strong>fspring <strong>of</strong> one larva are <strong>of</strong> <strong>the</strong><br />
same sex as <strong>the</strong> parent larva. These paedogenetic larvae do not develop a sterna! spatula<br />
but it reappears when pupation is imminent which confirms <strong>the</strong> function <strong>of</strong> this structure<br />
as a prerequisite <strong>to</strong> pupation. Pupal paedogenesis is also known in <strong>the</strong> <strong>British</strong><br />
species Henria psalliotae Wyatt, a pest <strong>of</strong> mushrooms. This reproductive pupa is called<br />
a hemipupa (Wyatt, I. J., 1961).<br />
Some Chironomidae appear <strong>to</strong> be paedogenetic but actually <strong>the</strong> adult female lays her<br />
eggs while she is still inside <strong>the</strong> pupal skin and is thus a pharate adult.<br />
In most Nema<strong>to</strong>cera and Brachycera <strong>the</strong> fully fed larvae form pupae but in<br />
Cyclorrhapha <strong>the</strong> last (third) instar larva forms a puparium from <strong>the</strong> last larval skin<br />
inside which it pupates. Stratiomyidae are exceptional in <strong>the</strong> Brachycera in not shedding<br />
<strong>the</strong> last larval skin when <strong>the</strong>y pupate but this appears <strong>to</strong> be a protective device unrelated<br />
<strong>to</strong> <strong>the</strong> cyclorrhaphous puparium. Some Nema<strong>to</strong>cera also pupate inside <strong>the</strong> last larval<br />
skin, e.g. Mayetiola and Chor<strong>to</strong>myia (Cecidomyiidae), some Sca<strong>to</strong>psidae, Pen<strong>the</strong>tria<br />
holosericea Meigen (non-<strong>British</strong>- Bibionidae) and <strong>the</strong> non-<strong>British</strong> Perissomatidae.<br />
Eggs<br />
The eggs <strong>of</strong> flies are generally oval or spindle-shaped, white or pale yellow and<br />
relatively featureless <strong>to</strong> <strong>the</strong> naked eye. Under <strong>the</strong> microscope many are seen <strong>to</strong> have a<br />
sculptured surface and some also have horns, stalks or a raised network <strong>of</strong> ridges. These<br />
surface features are largely respira<strong>to</strong>ry adaptations <strong>to</strong> prevent <strong>the</strong> eggs from drowning<br />
after rain and function as a respira<strong>to</strong>ry plastron which holds a thin film <strong>of</strong> air, as in<br />
many aquatic beetles and bugs (Hin<strong>to</strong>n, 1961 ). The ornamentation <strong>of</strong> an egg increases<br />
surface tension and may cause it <strong>to</strong> float, and <strong>the</strong> eggs <strong>of</strong> aquatic species may have more<br />
elaborate floatation devices (e.g. mosqui<strong>to</strong>es, figs 976-978). At <strong>the</strong> head end <strong>of</strong> <strong>the</strong> egg<br />
is <strong>the</strong> micropyle (fig. I 023), an opening for <strong>the</strong> admission <strong>of</strong> sperm.<br />
The largest number <strong>of</strong> eggs (c. 2,000) is produced by some aquatic flies whose larvae<br />
suffer enormous mortality from predation (e.g. Chironomus plumosus L.). Female<br />
blow-flies (Calliphora) lay about 300 eggs, usually in several small batches. The female<br />
housefly (Musca domestica L.) lays eggs singly in small or large batches at <strong>the</strong> rate <strong>of</strong><br />
I 00-150 per day up <strong>to</strong> I ,000. At <strong>the</strong> o<strong>the</strong>r extreme, Hippoboscidae and o<strong>the</strong>r Pupipara<br />
mature only one egg at a time, nurture it internally and eventually deposit a fully<br />
24
grown larva. Some families (e.g. Sarcophagidae) are normally viviparous and produce<br />
living first stage larvae, but when food is scarce even <strong>the</strong> bluebottle (Calliphora,<br />
Calliphoridae) will retain its eggs and produce living first stage larvae.<br />
Probably <strong>the</strong> eggs <strong>of</strong> <strong>the</strong> majority <strong>of</strong> Diptera are dropped in flight and roll in<strong>to</strong><br />
crevices in <strong>the</strong> ground but some are placed more deliberately and individually on a<br />
suitable medium in which <strong>the</strong> larvae can develop and this is especially so with parasitic<br />
species.<br />
Hin<strong>to</strong>n (1981) does not include Diptera in his tabulation <strong>of</strong> parental care in insects<br />
but A<strong>the</strong>rix (A<strong>the</strong>ricidae) exhibits something approaching this. The females practise<br />
communal egg-laying on leaves overhanging streams where <strong>the</strong>y <strong>the</strong>n die in masses like<br />
swarms <strong>of</strong> bees. The larvae hatch in <strong>the</strong> midst <strong>of</strong> <strong>the</strong> dead and dying females and drop<br />
in<strong>to</strong> <strong>the</strong> water. Females <strong>of</strong> <strong>the</strong> Nearctic tabanid Goniops chrysocoma Os ten Sacken stay<br />
with <strong>the</strong>ir eggs until <strong>the</strong>y hatch.<br />
There is no general work on <strong>the</strong> identification <strong>of</strong> <strong>the</strong> eggs <strong>of</strong>Diptera but illustrations<br />
<strong>of</strong> a wide range <strong>of</strong> eggs are given in this Handbook (figs 968-11 08). The major work <strong>of</strong><br />
Hin<strong>to</strong>n ( 1981) on insect eggs will be found useful, especially on microstructure (see also<br />
Salkeld, 1980).<br />
Since <strong>the</strong> egg is so neglected in descriptions it is worth remembering that this stage<br />
can <strong>of</strong>ten be obtained from live female adults whose identity can usually be established<br />
with certainty. Eggs may also be recovered from dry museum specimens when <strong>the</strong><br />
oviposi<strong>to</strong>r and abdomen are being macerated prior <strong>to</strong> dissection and study. The precise<br />
site <strong>of</strong> capture <strong>of</strong> <strong>the</strong> female, coupled with <strong>the</strong> structure <strong>of</strong> <strong>the</strong> egg and oviposi<strong>to</strong>r may<br />
also yield clues as <strong>to</strong> possible oviposition media in which development may take place<br />
(Smith, in Stubbs & Chandler, 1978: 247).<br />
Larvae<br />
The larvae <strong>of</strong> Diptera are very variable in appearance but may be distinguished from<br />
most o<strong>the</strong>r insect larvae by <strong>the</strong> lack <strong>of</strong> jointed thoracic legs, a ra<strong>the</strong>r slender form and<br />
usually active directional movement. A general study <strong>of</strong> larval morphology is provided<br />
by Teskey (in McAipine et al. 1981) which has proved invaluable in <strong>the</strong> preparation <strong>of</strong><br />
<strong>the</strong> following brief account. Some detailed studies <strong>of</strong> particular larval structures are<br />
cited in <strong>the</strong> appropriate sections below.<br />
Head<br />
The structure <strong>of</strong> <strong>the</strong> head varies considerably but three types may be recognised:<br />
eucephalic- with a well-developed head capsule and distinct, <strong>to</strong>o<strong>the</strong>d mandibles which<br />
move against each o<strong>the</strong>r in a horizontal or oblique plane (most Nema<strong>to</strong>cera, figs 2-4);<br />
hemicephalic-with <strong>the</strong> head capsule incomplete, partly retractile, and sickle-shaped<br />
mandibles which move in a vertical plane (Orthorrhapha-Brachycera, figs 166-167,<br />
170, 185); acepha/ic-<strong>the</strong> head capsule is fur<strong>the</strong>r reduced and retracted in<strong>to</strong> <strong>the</strong> thorax<br />
and forms, with <strong>the</strong> mouthparts, a cephalopharyngea/ skele<strong>to</strong>n (Cyclorrhapha, fig. 788).<br />
In <strong>the</strong> Cecidomyiidae <strong>the</strong> head capsule is almost absent except for <strong>the</strong> sterna/ spatula<br />
(sometimes referred <strong>to</strong> as <strong>the</strong> 'breastbone'), a peculiar ventral structure which is<br />
regarded as characteristic <strong>of</strong> <strong>the</strong> family though even this is absent in some species<br />
(figs. I 05-116). It is present only in <strong>the</strong> third ins tar (but not in paedogenetic forms) and<br />
is used for tunnelling in <strong>the</strong> soil.<br />
Some aquatic nema<strong>to</strong>cerous larvae (e.g. Simuliidae, Culicidae, some Chironomidae),<br />
in addition <strong>to</strong> <strong>the</strong> mandibles, have <strong>the</strong> labrum modified in<strong>to</strong> mouthbrushes<br />
(figs 43-44, br) which are used <strong>to</strong> sweep microplank<strong>to</strong>n <strong>to</strong>wards <strong>the</strong> mouth opening. In<br />
some aquatic non-predaceous Nema<strong>to</strong>cera (e.g. <strong>An</strong>isopodidae, Culicidae, Psychodidae,<br />
Ptychopteridae, Sca<strong>to</strong>psidae, Trichoceridae and some Tipulidae) <strong>the</strong> pharynx<br />
has a complex filtering apparatus for straining food particles from <strong>the</strong> water. Except<br />
for some aquatic larval Stratiomyidae this structure is absent in larvae <strong>of</strong> <strong>the</strong><br />
25
Orthorrhapha-Brachycera which are mostly predaceous. In <strong>the</strong>se Brachycera <strong>the</strong><br />
pharynx is sclerotized and in some families (e.g. Dolichopodidae, Empididae, Stratiomyidae)<br />
may also be fused with <strong>the</strong> internal skele<strong>to</strong>n <strong>of</strong> <strong>the</strong> head. In <strong>the</strong> Cyclorrhapha<br />
<strong>the</strong> head consists <strong>of</strong> an outer membranous segment and <strong>the</strong> internal cephalopharyngeal<br />
skele<strong>to</strong>n, a characteristic feature <strong>of</strong> <strong>the</strong> familiar maggot type <strong>of</strong> larva. The cephalopharyngeal<br />
skele<strong>to</strong>n is normally divisible in<strong>to</strong> three main parts: <strong>the</strong> basal or pharyngeal<br />
sclerite (ps); <strong>the</strong> intermediate or hypopharyngeal sclerite (hs) (ten<strong>to</strong>ropharyngeal) and<br />
<strong>the</strong> mouth hooks or mandibles (m) (fig. 788). The pharyngeal sclerite consists <strong>of</strong> a pair<br />
<strong>of</strong> roughly U-shaped sclerites on ei<strong>the</strong>r side <strong>of</strong> <strong>the</strong> pharynx. The two arms <strong>of</strong> each <strong>of</strong><br />
<strong>the</strong>se sclerites are called <strong>the</strong> dorsal (de) and ventral cornua (vc) (singular cornu) or<br />
'wings' and <strong>the</strong> latter are fused <strong>to</strong> <strong>the</strong> pharynx on each side. The pharyngeal sclerites<br />
may be joined by a bridge anterodorsally and a pair <strong>of</strong> slender paras<strong>to</strong>mal bars may<br />
project from <strong>the</strong> anterior margin above <strong>the</strong> hypos<strong>to</strong>mal sclerite (fig. 788, p). Below <strong>the</strong><br />
basal part <strong>of</strong> <strong>the</strong> mandible <strong>the</strong>re may be a dental sclerite (ds) and beneath (Muscidae,<br />
fig. 911, ob, ar) or between (Calliphoridae, fig. 788) <strong>the</strong> apices <strong>of</strong> <strong>the</strong> mandibles o<strong>the</strong>r<br />
accessory oral sclerites (os) may be present. Papers by Roberts (1969- 197la) should be<br />
consulted for a detailed discussion <strong>of</strong> <strong>the</strong> mouthparts <strong>of</strong> larval Diptera in relation <strong>to</strong><br />
feeding habits. A knowledge <strong>of</strong> <strong>the</strong> functional morphology <strong>of</strong> mouth parts may provide<br />
important clues as <strong>to</strong> whe<strong>the</strong>r or not an unidentified larva is <strong>the</strong> cause <strong>of</strong> primary<br />
damage <strong>to</strong> its host.<br />
The antennae (fig. 2) are usually close <strong>to</strong> <strong>the</strong> anterior mandibular articulations near<br />
<strong>the</strong> anterodorsal corners <strong>of</strong> <strong>the</strong> genae (cheeks). They may be very small (e.g.<br />
<strong>An</strong>isopodidae, Bibionidae, Ptychopteridae, some Psychodidae, many Myce<strong>to</strong>philidae)<br />
and are normally subdivided in<strong>to</strong> three divisions (though in some Chironomidae seven<br />
may be evident). In Chaoboridae <strong>the</strong> antennae have evolved in<strong>to</strong> prehensile structures<br />
with apical spines used for capturing prey (fig. 45, an).<br />
The eyes are also located on <strong>the</strong> genae. In <strong>the</strong> Nema<strong>to</strong>cera and orthorrhaphous<br />
Brachycera <strong>the</strong> eyes are usually simple (double in many Chironomidae) but in some<br />
Culicidae and Chaoboridae a compound eye is present, in front <strong>of</strong> <strong>the</strong> simple eye<br />
(ocellus). Little work has been done on <strong>the</strong> eyes <strong>of</strong>larval Diptera but Roberts (1970b) is<br />
useful. <strong>An</strong> egg-burster or hatching spine may be present on <strong>the</strong> head <strong>of</strong> <strong>the</strong> first instar<br />
larva <strong>of</strong> some Diptera (fig. 984) (see Edwards, F. W., 1919; Madwar, 1934; Hin<strong>to</strong>n,<br />
1981 ). Egg-bursters in Diptera require fur<strong>the</strong>r investigation (see Smith, 1955a).<br />
Body<br />
This is very variable in shape. Most Nema<strong>to</strong>cera and some Brachycera are slender<br />
and subcylindrical; Therevidae, Scenopinidae and some Ephydridae and Canacidae<br />
are spindle-shaped (fusiform); Cyclorrhapha and Xylophagidae are markedly<br />
narrowed anteriorly; Fanniidae, Lonchopteridae, Platypezidae, Stratiomyidae and<br />
Xylomyidae are dorsoventrally flattened; Chaoboridae and Culicidae have a swollen<br />
thorax; Simuliidae are swollen posteriorly while some Syrphidae and parasitic groups<br />
are generally s<strong>to</strong>ut; Conopidae are pear shaped and Microdon (Syrphidae) species are<br />
hemispherical.<br />
Nema<strong>to</strong>cera usually have 12 body segments, o<strong>the</strong>r Diptera 11. Three <strong>of</strong> <strong>the</strong>se<br />
segments comprise <strong>the</strong> thorax and <strong>the</strong> remainder <strong>the</strong> abdomen. Variation in <strong>the</strong><br />
numbers <strong>of</strong> abdominal segments nearly always involves a decrease in Nema<strong>to</strong>cera and<br />
an increase in o<strong>the</strong>r Diptera.<br />
In <strong>the</strong> Chaoboridae, Culicidae and Simuliidae <strong>the</strong> three thoracic segments are fused.<br />
In <strong>the</strong> <strong>An</strong>isopodidae and Psychodidae subdivision <strong>of</strong> <strong>the</strong> segments, or pseudosegmentation,<br />
occurs (figs 40, 70, 71). The Therevidae and Scenopinidae have 20 segmental<br />
divisions.<br />
The cuticle <strong>of</strong>Diptera larvae is usually only weakly sclerotized and non-pigmented, a<br />
condition probably explained by <strong>the</strong>ir mainly concealed mode <strong>of</strong> life in a humid terrestrial<br />
atmosphere or by <strong>the</strong> adoption <strong>of</strong> an aquatic or semi-aquatic environment. Larvae<br />
26
living in a drier pabulum have a <strong>to</strong>ugher cuticle <strong>to</strong> protect <strong>the</strong>m from abrasion and<br />
water loss (e.g. Asilidae, Xylophagidae). The larvae <strong>of</strong>Stratiomyidae and Xylomyidae<br />
have a <strong>to</strong>ugh cuticle <strong>of</strong> a shagreened appearance caused by <strong>the</strong> deposition <strong>of</strong> calcium<br />
carbonate, apparently an adaptation <strong>to</strong> ei<strong>the</strong>r periodic drought in normally moist<br />
terrestrial or temporary aquatic habitats (Hin<strong>to</strong>n, 1953), or for survival in acid<br />
conditions which cannot be <strong>to</strong>lerated by <strong>the</strong> larvae <strong>of</strong> o<strong>the</strong>r species not having this<br />
characteristic (McFadden, 1967). The larvae <strong>of</strong> some Pericoma species (Psychodidae)<br />
are covered in a layer <strong>of</strong> mud trapped by curved hairs on <strong>the</strong> dorsal surface.<br />
In colour <strong>the</strong> larvae <strong>of</strong> most Diptera are yellowish or whitish, probably reflecting<br />
<strong>the</strong>ir habit <strong>of</strong> usually living in concealment (e.g. in soil, dung, stems, wood, leaf mines,<br />
etc.). Larvae that live on <strong>the</strong> surface <strong>of</strong> plants may be greenish, especially if <strong>the</strong>y feed on<br />
<strong>the</strong> plants (e.g. Tipulidae, subfamily Cylindro<strong>to</strong>minae). The larvae <strong>of</strong> aphidophagous<br />
Syrphidae also live on <strong>the</strong> surface <strong>of</strong> plants, searching for <strong>the</strong>ir prey and are <strong>of</strong>ten quite<br />
brightly coloured. The red coloration <strong>of</strong> some larval Chironomidae is due <strong>to</strong> haemoglobin<br />
in <strong>the</strong> haemolymph. Similarly <strong>the</strong> brownish colour <strong>of</strong> some Tabanidae is due <strong>to</strong><br />
coloration <strong>of</strong> <strong>the</strong> haemolymph. Some Tipulidae have dense microtrichia on <strong>the</strong> cuticle<br />
which may give <strong>the</strong>m a grey or golden (some Limoniinae) appearance.<br />
In addition <strong>to</strong> hairs and bristles, which may be modified, <strong>the</strong> cuticle <strong>of</strong>Diptera larvae<br />
may have minute spicules, and scale-like, larger projections. These may be arranged<br />
in groups or patterns and thus be <strong>of</strong> use taxonomically. Tubercles or larger fleshy<br />
processes may also be present on <strong>the</strong> body segments, especially on <strong>the</strong> last segment<br />
where <strong>the</strong>y may form a series around <strong>the</strong> spiracles: <strong>the</strong>ir particular arrangement can be<br />
<strong>of</strong> use in identification.<br />
Some cuticular outgrowths may be used in locomotion. True jointed thoracic legs are<br />
never present in Diptera, but two types <strong>of</strong> false legs have evolved. Prolegs usually occur<br />
in pairs, ventrally, on <strong>the</strong> thoracic and anal segments, and sometimes on <strong>the</strong> abdominal<br />
segments (figs 20 I, 938). These pro legs bear one or more locomo<strong>to</strong>ry spinules at <strong>the</strong>ir<br />
apices (figs 152-156, 938, 945). Creeping welts are transverse swollen ridges usually<br />
found on <strong>the</strong> anterior ventral margins and sometimes <strong>the</strong> anterior dorsal margins <strong>of</strong> <strong>the</strong><br />
first seven body segments. These locomo<strong>to</strong>ry welts usually have transverse rows <strong>of</strong><br />
spinules, <strong>the</strong> microstructure and arrangement <strong>of</strong> which may have diagnostic importance<br />
(figs 529, 564). Larval creeping welts are commonest in Schizophora but also<br />
occur in some Nema<strong>to</strong>cera, and Brachycera-Orthorrhapha. Some non-<strong>British</strong> species<br />
(e.g. Blephariceridae and some Psychodidae) have developed suction discs which<br />
anchor <strong>the</strong>m in swiftly flowing water. O<strong>the</strong>r backwardly directed bristles or spines or<br />
cuticular ridges may also assist in locomotion. Roberts (1971b) discusses locomotion<br />
by apodous cyclorrhaphan maggots.<br />
Respira<strong>to</strong>ry System<br />
There are seven main types <strong>of</strong> respira<strong>to</strong>ry system in <strong>the</strong> larvae <strong>of</strong>Diptera (fig. 1) and<br />
<strong>the</strong>se may vary not only between taxa or habitat types occupied but also between<br />
instars <strong>of</strong> <strong>the</strong> same species. The systems described below refer <strong>to</strong> <strong>the</strong> final instar<br />
larvae. Detailed discussions <strong>of</strong> respira<strong>to</strong>ry systems and <strong>the</strong>ir systematic importance are<br />
provided by Keilin (1944) and Whitten (1959).<br />
The holopneustic type is <strong>the</strong> basic system in which <strong>10</strong> pairs <strong>of</strong> spiracles are present,<br />
located on <strong>the</strong> prothorax, metathorax and eight <strong>of</strong> <strong>the</strong> abdominal segments (not on<br />
segment 9 in <strong>British</strong> genera). This condition occurs only in Bibionidae (figs 1, 74) and<br />
<strong>the</strong> non-<strong>British</strong> Pachyneuridae and Cramp<strong>to</strong>nomyidae. In larval descriptions <strong>the</strong><br />
prothoracic spiracles are called <strong>the</strong> anterior spiracles and those on <strong>the</strong> anal segment<br />
are termed <strong>the</strong> posterior spiracles. Both pairs are larger than <strong>the</strong> o<strong>the</strong>r, abdominal,<br />
spiracles.<br />
In <strong>the</strong> peripneustic system <strong>the</strong> metathoracic spiracles are absent and this is characteristic<br />
<strong>of</strong>Cecidomyiidae, Myce<strong>to</strong>philidae (sub-family Di<strong>to</strong>myiinae), Sca<strong>to</strong>psidae and <strong>the</strong><br />
non-<strong>British</strong> Synneuridae and Pachyneuridae o<strong>the</strong>r than Pachyneura.<br />
27
Posterior spiracles are lacking in <strong>the</strong> hemipneustic system which is represented by<br />
Sciaridae and most Myce<strong>to</strong>philidae.<br />
The amphipneustic system has only <strong>the</strong> anterior and posterior pairs <strong>of</strong> spiracles. It<br />
is <strong>the</strong> commonest system in Diptera and typical <strong>of</strong> <strong>An</strong>isopodidae, Psychodidae,<br />
Thaumaleidae, Trichoceridae (and <strong>the</strong> non-<strong>British</strong> Axymiidae and Tanyderidae) in <strong>the</strong><br />
Nema<strong>to</strong>cera and most o<strong>the</strong>r Diptera.<br />
In <strong>the</strong> propneustic condition only anterior spiracles are present. This respira<strong>to</strong>ry<br />
system occurs only in some Myce<strong>to</strong>philidae (Diadocidia species and some Sciophilinae).<br />
The metapneustic system in which only <strong>the</strong> posterior spiracles remain is found in<br />
some larvae living in aquatic or semi-aquatic conditions in <strong>the</strong> families Culicidae,<br />
Dixidae, Ptychopteridae, and Tipulidae in <strong>the</strong> Nema<strong>to</strong>cera, and <strong>the</strong> Tabanidae in <strong>the</strong><br />
Brachycera. The Tabanidae do have anterior spiracles but <strong>the</strong>y are extruded just before<br />
pupation and it is not known if <strong>the</strong>y are functional.<br />
When spiracles are completely absent (although tracheae are always welldeveloped)<br />
<strong>the</strong> condition is said <strong>to</strong> be apneustic. Such larvae are aquatic (except<br />
Keroplatinae, Myce<strong>to</strong>philidae) and include <strong>the</strong> Cera<strong>to</strong>pogonidae, some Chaoboridae<br />
(Chaoborus}, most Chironomidae, Simuliidae (and <strong>the</strong> non-<strong>British</strong> Blephariceridae,<br />
Deuterophlebiidae and Nymphomyiidae) in <strong>the</strong> Nema<strong>to</strong>cera; A<strong>the</strong>ricidae and<br />
Empididae in <strong>the</strong> Brachycera; and some representatives <strong>of</strong> o<strong>the</strong>r families.<br />
The respira<strong>to</strong>ry system may differ between instars, e.g. Myce<strong>to</strong>philidae and<br />
Sciaridae are metapneustic in <strong>the</strong> first ins tar, propneustic in <strong>the</strong> second and third ins tars<br />
and hemipneustic in <strong>the</strong> fourth (final) ins tar. In Cyclorrhapha <strong>the</strong> first ins tar larvae are<br />
metapneustic and <strong>the</strong> second and third (final) instar are usually amphipneustic.<br />
Spiracles vary in structure and Keilin (1944) recognised three types (in final stage<br />
larvae). In <strong>the</strong> first type <strong>the</strong> spiracular opening through which <strong>the</strong> trachea <strong>of</strong> <strong>the</strong><br />
previous instar is withdrawn becomes <strong>the</strong> spiracular opening <strong>of</strong> <strong>the</strong> following instar<br />
(e.g. Culicidae, Tabanidae). In <strong>the</strong> second type <strong>the</strong> spiracular opening is closed and in<br />
<strong>the</strong> following instar forms a scar around which perforated oval or linear air intakes<br />
open (e.g. most Nema<strong>to</strong>cera and Brachycera). In type three <strong>the</strong> spiracular opening <strong>of</strong><br />
<strong>the</strong> second instar forms a scar near <strong>the</strong> inner margin <strong>of</strong> <strong>the</strong> spiracular plate and <strong>the</strong><br />
number <strong>of</strong> air intakes opening around <strong>the</strong> scar is usually reduced (e.g. Cyclorrhapha).<br />
A very useful 'spot character' <strong>of</strong> <strong>the</strong> larvae <strong>of</strong> most Cyclorrhapha is that in <strong>the</strong> three<br />
instars each posterior spiracle has one, two or three respira<strong>to</strong>ry slits respectively (figs<br />
786, 790). The spiracular plate upon which <strong>the</strong> posterior spiracles are situated has a<br />
margin or peritreme. The arrangement and shape <strong>of</strong> <strong>the</strong> spiracular slits and <strong>the</strong> state <strong>of</strong><br />
<strong>the</strong> peritreme (complete or incomplete) provide valuable diagnostic characters in <strong>the</strong><br />
Cyclorrhapha.<br />
In <strong>the</strong> majority <strong>of</strong> <strong>the</strong> Nema<strong>to</strong>cera <strong>the</strong> posterior spiracles are usually sessile but in<br />
<strong>the</strong> Brachycera and Cyclorrhapha <strong>the</strong>y are raised above <strong>the</strong> body surface on short<br />
processes. Larvae living in liquid or semi-liquid media may have <strong>the</strong> spiracles each<br />
situated at <strong>the</strong> end <strong>of</strong> two short processes (Sca<strong>to</strong>psidae) or <strong>to</strong>ge<strong>the</strong>r at <strong>the</strong> end <strong>of</strong> a<br />
relatively short respira<strong>to</strong>ry siphon (e.g. Psychodidae, Culicidae, Stratiomyidae,<br />
Tabanidae and Ephydridae) or at <strong>the</strong> end <strong>of</strong> a long and sometimes retractable siphon<br />
(e.g. Ptychopteridae; Eristalis and some o<strong>the</strong>r Syrphidae; Aulacigastridae).<br />
Some Stratiomyidae exhibit ano<strong>the</strong>r adaptation <strong>to</strong> an aquatic existence: <strong>the</strong>y have<br />
<strong>the</strong> posterior spiracles enclosed in a small chamber (atrium}, <strong>the</strong> opposing edges <strong>of</strong><br />
which are fringed with hairs which enclose a bubble <strong>of</strong> air when <strong>the</strong> larva submerges<br />
(fig. 135). The same effect can be achieved by <strong>the</strong> anal lobes especially when <strong>the</strong>y are<br />
fringed with hairs (e.g. Tipulidae (fig. 27) and Dolichopodidae (figs 208- 209). The<br />
spiracular pit in Sarcophaga (fig. 781) may have a similar function.<br />
The posterior spiracles <strong>of</strong> Chrysogaster (Syrphidae, fig . 258) and Notiphila<br />
(Ephydridae, figs 615-616) are modified in<strong>to</strong> a respira<strong>to</strong>ry spine used <strong>to</strong> pierce and<br />
draw air from tissues <strong>of</strong> aquatic plants. Structurally similar spines are found in some<br />
Tabanidae but are not situated at <strong>the</strong> point <strong>of</strong> air intake, and spiracular spines <strong>of</strong><br />
28
unknown function are present in some Acalyptrates (e.g. Clusiidae, Lonchaeidae,<br />
Micropezidae and Psilidae).<br />
In <strong>the</strong> Cyclorrhapha <strong>the</strong> anterior spiracles <strong>of</strong>ten have several openings, <strong>the</strong> number<br />
and arrangement <strong>of</strong> which may provide characters <strong>of</strong> taxonomic value.<br />
In many aquatic or semi-aquatic larvae <strong>of</strong> Nema<strong>to</strong>cera, anal papillae may be<br />
present, near or within <strong>the</strong> anus on <strong>the</strong> last abdominal segment. These appear <strong>to</strong> have<br />
mainly an osmoregula<strong>to</strong>ry function (Wigglesworth, 1938; Brindle, 1952) but when well<br />
tracheated have been called tracheal gills, though <strong>the</strong> respira<strong>to</strong>ry function is in doubt.<br />
A list <strong>of</strong> <strong>the</strong> late Pr<strong>of</strong>essor H. E. Hin<strong>to</strong>n's important contributions <strong>to</strong> our understanding<br />
<strong>of</strong> respira<strong>to</strong>ry systems in Diptera (and o<strong>the</strong>r insects) is included in <strong>the</strong><br />
bibliography <strong>of</strong> his publications at <strong>the</strong> end <strong>of</strong> vol. 3 <strong>of</strong> his work on insect eggs (Hin<strong>to</strong>n,<br />
1981 ), and a few are cited at appropriate places in this Handbook.<br />
Useful works on <strong>the</strong> identification <strong>of</strong> larvae are cited in <strong>the</strong> <strong>introduction</strong> and under<br />
each family below. A major work on <strong>the</strong> larvae <strong>of</strong>Cyclorrhapha has been published as<br />
this work goes <strong>to</strong> press (Ferrar, 1987).<br />
Pupae<br />
The pupae <strong>of</strong> Diptera are <strong>of</strong> two main types.<br />
In obtect pupae (figs 1<strong>10</strong>9-1148) <strong>the</strong> head appendages, wings and legs are visible and<br />
lie in sheaths attached <strong>to</strong> <strong>the</strong> surface <strong>of</strong> <strong>the</strong> body (e.g. Nema<strong>to</strong>cera and Brachycera).<br />
Obtect pupae are mostly immobile although <strong>the</strong>y can move by wriggling <strong>the</strong> abdomen.<br />
Pupae <strong>of</strong> this type may be enclosed in a silken cocoon covered with soil or o<strong>the</strong>r<br />
particles. Pupae <strong>of</strong> aquatic species may be able <strong>to</strong> swim actively (e.g. Culicidae,<br />
Chironomidae, Chaoboridae). The abdominal segments and head <strong>of</strong>ten have bristles<br />
or hairs which help <strong>the</strong> pupa move through or break out <strong>of</strong> <strong>the</strong> soil, wood or o<strong>the</strong>r<br />
medium in which it has developed. The adult emerges through a rectangular or T<br />
shaped rupture (from which <strong>the</strong> name Orthorrhapha is derived: Gr. ortho-straight,<br />
raphe-suture).<br />
The pupae <strong>of</strong> Forcipomyia and Atrichopogon retain <strong>the</strong> larval skin over <strong>the</strong> apical<br />
half <strong>of</strong> <strong>the</strong> body (fig. 1120). Only a few o<strong>the</strong>r Orthorrhapha do this, e.g. Stratiomyidae<br />
(entire cover), Sca<strong>to</strong>psidae (thorax free), Dactylolabis and Cylindro<strong>to</strong>ma (Tipulidae)<br />
(partially covered) and some Chironomidae that pupate inside a tube or case do not<br />
free <strong>the</strong>mselves from a larval skin until <strong>the</strong>y wriggle out <strong>to</strong> escape <strong>to</strong> <strong>the</strong> water surface.<br />
The second type <strong>of</strong> pupa is found in Cyclorrhapha and is exarate, i.e. <strong>the</strong> appendages<br />
are free. However <strong>the</strong> pupa is rarely visible as pupation takes place inside <strong>the</strong> last larval<br />
skin or puparium (figs 792-793), 1149-1286) which is rounded, hardened and tanned. In<br />
some families <strong>the</strong> pupa or puparium may be concealed inside a cocoon made by <strong>the</strong><br />
larva before pupariation (figs 1142, 1145, 1156, 1272). The adult emerges by inflating a<br />
large sac on its head, <strong>the</strong> ptilinum, which lifts a circular cap <strong>of</strong>f <strong>the</strong> puparium (from<br />
which <strong>the</strong> name Cyclorrhapha is derived: Gr. cyclos-round; raphe-suture). The<br />
puparial cap splits in two, and one half contains <strong>the</strong> mouthparts <strong>of</strong> <strong>the</strong> final ins tar larva.<br />
The mouth parts, in conjunction with <strong>the</strong> posterior spiracles retained on <strong>the</strong> surface <strong>of</strong><br />
<strong>the</strong> puparium, may render identification <strong>of</strong>puparia possible. This can be invaluable in<br />
certain types <strong>of</strong> investigation especially forensic and archaeological work (Smith,<br />
1986b), when puparia are frequently <strong>the</strong> only material available for study.<br />
Most pupae brea<strong>the</strong> entirely through <strong>the</strong> thoracic spiracles which are sometimes<br />
produced in<strong>to</strong> pupal horns (figs 11<strong>10</strong>--1122, 1143-1145). In some aquatic species such<br />
pupal horns may be sharp and capable <strong>of</strong> piercing plant tissues in order <strong>to</strong> obtain<br />
oxygen.<br />
The pupal stage is <strong>of</strong>ten neglected in descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> unless it<br />
is <strong>the</strong> stage most frequently found (e.g. Asilidae, Sphaeroceridae) or <strong>of</strong> particular<br />
taxonomic or economic (especially medical) importance (e.g. Culicidae, Simuliidae,<br />
Chironomidae).<br />
29
To facilitate approximate identification a range <strong>of</strong> pupae and puparia is illustrated<br />
(figs 1<strong>10</strong>9-1286) for most families. Brauns (1954b) is useful for terrestrial species and<br />
o<strong>the</strong>r works dealing substantially with pupae are indicated under <strong>the</strong> appropriate<br />
families (see Ferrar, 1987 for Cyclorrhapha).<br />
Classification, nomenclature and use <strong>of</strong> keys<br />
As far possible <strong>the</strong> sequence <strong>of</strong> families, subfamilies, tribes and genera follows that <strong>of</strong><br />
<strong>the</strong> Diptera Check List (Smith et al., 1976) for <strong>the</strong> convenience <strong>of</strong> users <strong>of</strong> <strong>the</strong> Handbook<br />
series. Sometimes <strong>the</strong> only workable keys <strong>to</strong> larvae are based on a classification different<br />
from that now used for <strong>the</strong> adults (e.g. Syrphidae) and in such cases explana<strong>to</strong>ry<br />
comments are given. Where recent taxonomic research on adults has led <strong>to</strong> changes in<br />
<strong>the</strong> nomenclature, status, or generic placing <strong>of</strong> well-known species (such as pests)<br />
adequate synonymy is cited under <strong>the</strong> appropriate family treatment. Colloquial names<br />
for pest species, plant crops and o<strong>the</strong>r familiar plants and animals are given in addition<br />
<strong>to</strong> <strong>the</strong>ir scientific names at <strong>the</strong> first (or most important) citation.<br />
In using <strong>the</strong> keys it should constantly be borne in mind that many genera, species and<br />
even some families are unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and <strong>the</strong>refore <strong>the</strong> full range <strong>of</strong><br />
morphological variety is unknown. <strong>Part</strong>icularly in <strong>the</strong> Acalyptratae couplets may lead<br />
<strong>to</strong> a choice <strong>of</strong> families and several families run through <strong>to</strong> more than one place in <strong>the</strong><br />
keys. It is intended that identifications should be attempted by combining use <strong>of</strong> <strong>the</strong><br />
keys with a consideration <strong>of</strong> habitat and pabulum (using <strong>the</strong> introduc<strong>to</strong>ry sections in<br />
combination with <strong>the</strong> index) in conjunction with <strong>the</strong> figures. In this way it should be<br />
possible <strong>to</strong> identify many <strong>of</strong> <strong>the</strong> Diptera most commonly found as larvae or pupae,<br />
including all species <strong>of</strong> economic, agricultural, medical and veterinary importance. As<br />
far as possible I have aimed <strong>to</strong> include all those Diptera whose <strong>immature</strong> <strong>stages</strong> are<br />
regularly submitted <strong>to</strong> en<strong>to</strong>mologists as 'economic enquiries'.<br />
Notes on <strong>the</strong> illustrations<br />
'As far as I am a judge, nothing would recommend en<strong>to</strong>mology more than<br />
some neat plates that should well express <strong>the</strong> generic distinctions <strong>of</strong> insects<br />
according <strong>to</strong> Linnaeus; for I am well assured that many people would study<br />
insects, could <strong>the</strong>y set out with a more adequate notion <strong>of</strong> those distinctions<br />
than can be conveyed at first by words alone.'<br />
Gilbert White, 1771<br />
The illustrations are meant <strong>to</strong> supplement <strong>the</strong> text with <strong>the</strong> following aims:<br />
(I) facilitating identification as far as family and tribe in conjunction with <strong>the</strong> keys;<br />
(2) <strong>to</strong> show <strong>the</strong> range <strong>of</strong> form and <strong>to</strong> illustrate particular environmental adaptations;<br />
(3) <strong>to</strong> facilitate <strong>the</strong> identification <strong>to</strong> genus and species <strong>of</strong> selected Diptera such as pest<br />
species, well known 'text-book' species and common readily encountered species <strong>of</strong><br />
value for teaching or demonstration, or frequently found during faunistic surveys.<br />
Attempts at identification by perusal <strong>of</strong> <strong>the</strong> whole insect figures alone should be<br />
practised with extreme caution making particular scrutiny <strong>of</strong> structural details and<br />
scanning all <strong>the</strong> figures. Convergence through adaptations <strong>to</strong> similar environmental<br />
requirements may result in remarkably similar superficial appearances between several<br />
taxonomically widely separated species, e.g. compare figs <strong>10</strong>27 and <strong>10</strong>59; figs 239 and<br />
903; figs 20 I, 617 and 938 etc.<br />
Allowance should be made for some variation in <strong>the</strong> structures illustrated, particularly<br />
in <strong>the</strong> pigmentation <strong>of</strong> <strong>the</strong> cephalopharyngeal skele<strong>to</strong>n. Apart from some<br />
expected variation within a species <strong>the</strong>re is evidence <strong>of</strong> geographic variation. Published<br />
30
figures <strong>of</strong> <strong>the</strong> same species by different authors always show some slight differences.<br />
Skidmore ( 1985) frequently illustrates more than one example <strong>of</strong> mouth parts in species<br />
<strong>of</strong> Muscidae and <strong>the</strong> differences illustrate this point. Mandibular wear may occur in<br />
older larvae <strong>of</strong> a given instar (<strong>the</strong> sterna) spatula <strong>of</strong> Cecidomyiidae may also become<br />
worn). The number <strong>of</strong> rays on an anterior spiracle may differ by one or two, even on<br />
each side <strong>of</strong> <strong>the</strong> same larva. The length <strong>of</strong> <strong>the</strong> caudal lobes may vary within a species in<br />
response <strong>to</strong> <strong>the</strong> amount <strong>of</strong> moisture in <strong>the</strong> habitat (see under Sca<strong>to</strong>psidae).<br />
However, by careful use <strong>of</strong> keys, illustrations, text (introduc<strong>to</strong>ry and under <strong>the</strong><br />
family) and index <strong>the</strong> author believes it should be possible <strong>to</strong> identify nearly all Diptera<br />
larvae <strong>to</strong> family, most <strong>to</strong> genus and a large number <strong>to</strong> species, including all <strong>of</strong> <strong>the</strong><br />
so-called 'pest' species.<br />
To save space in <strong>the</strong> figure legends (and <strong>to</strong> avoid irksome abbreviations) <strong>the</strong> cephalopharyngeal<br />
skele<strong>to</strong>n is referred <strong>to</strong> as 'mouthparts' throughout and acknowledgements<br />
<strong>to</strong> published sources <strong>of</strong> many <strong>of</strong> <strong>the</strong> figures are given at <strong>the</strong> beginning <strong>of</strong> <strong>the</strong> book.<br />
31
32<br />
Key <strong>to</strong> suborders <strong>of</strong> <strong>British</strong> Diptera larvae
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Nema<strong>to</strong>cera<br />
Head greatly reduced and lightly sclerotized; mouthparts indistinct but a variously shaped,<br />
prothoracic, 'sterna! spatula' is usually present ventrally (figs <strong>10</strong>3- 116). Larvae small,<br />
inactive; pink, red, orange, yellow or white in colour. Respira<strong>to</strong>ry system peripneustic or<br />
apneustic. Frequently inhabiting plant galls but also under bark where larvae may occur in<br />
masses, also found in soil or humus (figs <strong>10</strong>1- 122) Cecidomyiidae (p. 52)<br />
Head usually distinct and forming a sclerotized capsule; without a sterna! spatula;<br />
respira<strong>to</strong>ry system variable 2<br />
2 Head capsule complete or incomplete and reduced <strong>to</strong> a number <strong>of</strong> longitudinal sclerotized<br />
rods but always retractile, <strong>to</strong> some degree, in<strong>to</strong> <strong>the</strong> thorax. Larvae usually medium <strong>to</strong> large<br />
in size, elongate; active (carnivorous species) or slow moving (herbivores); yellowishwhitish<br />
or dark brown <strong>to</strong> greyish in colour; respira<strong>to</strong>ry system usually metapneustic, with<br />
one pair <strong>of</strong> spiracles on <strong>the</strong> last segment frequently surrounded by fleshy lobes (figs <strong>10</strong>-29)<br />
or rarely apneustic (e.g. <strong>the</strong> aquatic <strong>An</strong><strong>to</strong>cha); prolegs sometimes present (Pediciini)<br />
usually found in marshy soils but also in drier soils, in mosses, decayed or dead wood and<br />
sometimes aquatic (figs 9- 29) . Tipulidae (p. 35)<br />
Head capsule complete and non-retractile in<strong>to</strong> <strong>the</strong> thorax; respira<strong>to</strong>ry system usually<br />
o<strong>the</strong>rwise 3<br />
3 Abdomen long and slender with posterior margins <strong>of</strong> segments swollen and with a long<br />
retractile respira<strong>to</strong>ry tube on <strong>the</strong> anal segment, terminating in small unbranched anal<br />
papillae; in saturated marshy soils and mud (fig. 41) Ptychopteridae (p. 38)<br />
Abdomen not so long and slender; any respira<strong>to</strong>ry tube much shorter, thicker, and nonretractile<br />
4<br />
4 Prolegs present on at least one body segment 5<br />
Prolegs absent 9<br />
5 Paired pro legs on first two abdominal segments only; dorsum <strong>of</strong> abdomen may have rings (r)<br />
<strong>of</strong> setae on some segments; anal segment tapering posteriorly and with a dorsoventrally<br />
flattened leaf-like process (p) on each side. Very active larvae usually curved in a U-shape<br />
in life; on surface or slower moving water, on wet rocks or mosses (fig. 42) .<br />
Dixidae (p. 39)<br />
Pro legs ei<strong>the</strong>r on thoracic segments or on posterior abdominal segments 6<br />
6 Head capsule with a pair <strong>of</strong> conspicuous, folding, dorsolateral mouth-brushes (labral fans)<br />
(fig. 69); abdomen conspicuously swollen distally; anal segment with circlet <strong>of</strong> minute<br />
hooked spines. Larvae sessile, usually anchored by <strong>the</strong> circlet <strong>of</strong> spines on <strong>the</strong> anal segment<br />
<strong>to</strong> pad <strong>of</strong> silk attached <strong>to</strong> s<strong>to</strong>nes, weed, etc. in flowing water (figs 68-69) .<br />
Simuliidae (p. 45)<br />
Head capsule without such conspicuous mouth-brushes (Culicidae have hair tufts each side<br />
<strong>of</strong> <strong>the</strong> labrum); abdomen not conspicuously swollen distally; not sessile . 7<br />
7 Paired prolegs present on <strong>the</strong> first thoracic segment and on <strong>the</strong> last abdominal segment, with<br />
(figs 65- 66) or without (fig. 67) (except some Orthocladiinae, see below) additional gills.<br />
Larvae active, aquatic or terrestrial; aquatic larvae frequently swim with a looping<br />
movement (figs 53-67) . Chironomidae (p. 43)<br />
Pro legs on thoracic segments only, and not paired but fused and appearing bifurcated 8<br />
8 Body smooth, almost parallel-sided; without prominent setae but with sclerotized darker<br />
dorsal areas; respira<strong>to</strong>ry system amphipneustic with <strong>the</strong> anterior spiracles on a short stalk<br />
and <strong>the</strong> posterior spiracles opening in<strong>to</strong> a transverse cleft between fingerlike processes on<br />
<strong>the</strong> 8th abdominal segment. On rocks, mosses, etc., washed by a film <strong>of</strong> water<br />
(hygropetricous); larval movement active and sinuous (fig. 47) . Thaumaleidae (p. 41)<br />
Body not smooth, less parallel-sided with conspicuous setae which may be long and filiform<br />
(figs 49) or spatulate (fig 48). Larvae slow; found in moist habitats, in soil, under bark, with<br />
water drops <strong>of</strong>ten adhering <strong>to</strong> <strong>the</strong> setae (figs 48-49) . Cera<strong>to</strong>pogonidae (part) (p. 41)<br />
9 Body slender (almost thread-like) with uniform bead-like segments and conspicuous setae<br />
present only on terminal abdominal segment. Movement active, sinuous; in wet mosses<br />
(e.g. Sphagnum), sand <strong>of</strong> river banks or sometimes freely aquatic (figs 50-52)<br />
Cera<strong>to</strong>pogonidae (part) (p. 41)<br />
Body not so slender or smooth . <strong>10</strong><br />
<strong>10</strong> Thoracic segments fused and wider than <strong>the</strong> rest <strong>of</strong> <strong>the</strong> body 11<br />
Thoracic segments not fused nor distinctly wider than <strong>the</strong> rest <strong>of</strong> <strong>the</strong> body 12<br />
33
ll <strong>An</strong>tennae (an) prehensile (adapted for holding or seizing), with long strong apical spines;<br />
without distinct mouth-brushes; thorax and 7th abdominal segment with paired pigmented<br />
air sacs (as). Larvae active, but hold a horizontal position in <strong>the</strong> water betwee n<br />
movements; in still or slow moving water (figs 45-46) . Chaoboridae (p. 39)<br />
<strong>An</strong>tennae not prehensile, without strong apical spines; with distinct mouth-brushes, one on<br />
each side <strong>of</strong> labrum (figs 43-44, br); no pigmented air sacs. Larva active but usually hold<br />
an oblique (fig. 44) (horizontal in <strong>An</strong>opheles fig. 43) position at <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water<br />
between movements; in still or slow moving water (figs 43-44) . Culicidae (p. 39)<br />
12 Body with short tubercles or spinous processes on all segments; anal segment with longer<br />
processes; respira<strong>to</strong>ry system holopneustic with <strong>10</strong> pairs <strong>of</strong> spiracles. Larvae inactive,<br />
characteristically curved in life; usually communal in decaying organic matter, rich soil,<br />
plant roots, dung, etc. (figs 74-78) Bibionidae (p. 47)<br />
Body o<strong>the</strong>rwise; less than <strong>10</strong> pairs <strong>of</strong>spiracles present . 13<br />
13 The two posterior spiracles each on <strong>the</strong> tip <strong>of</strong> a cylindrical process (except Eclaetia fig . lOO).<br />
Larvae usually dark or at least brown or yellow and ra<strong>the</strong>r hairy; very inactive; in dung,<br />
soil or in drier habitats with decaying material, e.g. birds' nests, tree holes (figs 92 ~ <strong>10</strong>0)<br />
Sca<strong>to</strong>psidae (p. 50)<br />
The two posterior spiracles not on such processes 14<br />
14 Last abdominal segment usually with a short non-retractile respira<strong>to</strong>ry tube (figs 30, 32) or<br />
tergites <strong>of</strong> body with numerous distinct transverse sclerotized plates (figs 31 ~ 39) (except<br />
Trichomyia, fig 40); sometimes body with numerous processes. Larvae ra<strong>the</strong>r inactive, in<br />
decaying organic matter in aquatic or semi-aquatic situations (figs 30-40)<br />
Psychodidae (p. 37)<br />
Last abdominal segment without such a tube; abdomen without distinct transverse<br />
sclerotized plates . 15<br />
15 Abdominal segments subdivided superficially in<strong>to</strong> 3 more or less equal parts (fig. 5);<br />
posterior spiracles surrounded by four fleshy lobes (figs 6-8). Larva whitish <strong>to</strong> brownish,<br />
slow moving; in drier dung, soil, or in moist habitats with decaying organic material<br />
(figs 2~ 8) Trichoceridae (p. 35)<br />
Abdominal segments not subdivided superficially in<strong>to</strong> three more or less equal parts . 16<br />
16 Abdominal segments 2-6 divided superficially in<strong>to</strong> two unequal parts (figs 70-71); posterior<br />
spiracles surrounded by 5 reduced lobes (figs 72~73). Larvae slender, active; in moist<br />
habitats where decaying organic material occurs, in deliquescent fungi, or in drier habitats<br />
such as dung, soil, old plant stems, etc. (figs 70-73) . <strong>An</strong>isopodidae (p. 46)<br />
Abdominal segments not so divided; sometimes superficially ringed but in this case <strong>the</strong> rings<br />
are narrow, darker and equal; anal segment without lobes. Larvae usually white with<br />
contrasting darker, <strong>of</strong>ten black head capsule . 17<br />
17 Head capsule ventrally with epicranial plates meeting only at one point (or not meeting,<br />
Diadocidiinae) so that <strong>the</strong> posterior ten<strong>to</strong>rial bridge is absent (fig. 86); or if <strong>the</strong> bridge is<br />
partly formed <strong>the</strong> abdominal creeping welts have sclerotized spicules. Larvae usually<br />
white, sometimes dull yellow <strong>to</strong> bright green; usually in fungi , under bark, in silken webs in<br />
holes in old tree trunks, in soil, decaying plants, roots etc . Myce<strong>to</strong>philidae (p. 47)<br />
Head capsule ventrally meeting at two points, <strong>the</strong> posterior ten<strong>to</strong>rial bridge (ptb) being<br />
complete (figs 87~88) or nearly complete; abdominal creeping welts without sclerotized<br />
spicules. Larvae white with black head capsule; in decaying plant material, animal<br />
excrement, rotten wood attacked by fungus, some in old nests <strong>of</strong> mammals and birds,<br />
sometimes mass movements in large columns . Sciaridae (p. 50)<br />
34
Notes on families <strong>of</strong> Nema<strong>to</strong>cera<br />
Trichoceridae<br />
(Figs: larvae 2- 8, egg 968, pupa 1<strong>10</strong>9)<br />
The family Trichoceridae (winter-gnats) contains about 112 species in 5 genera <strong>of</strong><br />
which 9 species in 2 genera are known in <strong>the</strong> larval <strong>stages</strong>. In Britain 2 genera are<br />
represented: Diazosoma hirtipenne Siebke which is unknown in <strong>the</strong> larval stage and<br />
Trichocera with 9 species <strong>of</strong> which 6 are known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Trichocerid larvae are cylindrical and a little depressed, yellowish white <strong>to</strong> reddish<br />
brown in colour and amphipneustic. The cuticle is covered with fine pubescence. The<br />
posterior spiracles are surrounded by four fleshy lobes. In T. salta<strong>to</strong>r (Harris) <strong>the</strong><br />
dorso-lateral lobes are pigmented right up <strong>to</strong> <strong>the</strong> spiracle (fig. 7), in T. hiemalis (De<br />
Geer) <strong>the</strong> pigmentation ends short <strong>of</strong> <strong>the</strong> spiracle (fig. 6) and in <strong>the</strong> remaining species<br />
<strong>the</strong> lobes are unpigmented (fig. 8).<br />
The larvae are saprophagous and feed on decaying organic material. They have been<br />
found in pota<strong>to</strong>es, turnips and mangold-wurzels which have partly decayed when<br />
s<strong>to</strong>red in pits during <strong>the</strong> winter. T. salta<strong>to</strong>r and T. regelationis (L.) occur in dung;<br />
T. annulata (Meigen) and T. salta<strong>to</strong>r have been recorded from human corpses and<br />
probably occur on o<strong>the</strong>r carrion during <strong>the</strong> winter; I have received T. salta<strong>to</strong>r from a<br />
bottle <strong>of</strong> milk. Larvae also occur in fresh and decomposing fungi.<br />
The morphology <strong>of</strong> <strong>the</strong> larva and pupa <strong>of</strong>T. hiemalis is described by Keilin & Tate<br />
( 1940). Keys <strong>to</strong> species are provided by Brindle (19621), C. Dahl ( 1973) and Smith<br />
(1986b).<br />
Tipulidae<br />
(Figs: larvae 9- 29, eggs 969- 973, pupae Ill Q-1111 , emergence <strong>of</strong> adult 1327)<br />
The Tipulidae (crane-flies) is <strong>the</strong> largest family <strong>of</strong>Diptera (s~ also Tachinidae) with<br />
about 13,500 described species <strong>of</strong> which 303 species, representing 41 genera, occur in<br />
<strong>the</strong> <strong>British</strong> Isles. Of <strong>the</strong> <strong>British</strong> fauna about 150 species, representing 39 genera, are<br />
known in <strong>the</strong> larval stage. Larvae <strong>of</strong> <strong>the</strong> subfamilies Tipulinae and Cylindro<strong>to</strong>minae<br />
are fairly well known but in <strong>the</strong> Limoniinae many species are unknown in <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong>. Larvae <strong>of</strong> <strong>the</strong> three sub-families may be separated as follows (after Brindle,<br />
1967).<br />
Posterior spiracular disc with six lobes which are flattened or cylindrical, sometimes reduced<br />
in size and rarely entirely sclerotized (figs <strong>10</strong>-14) . . . . . . . . . Tipulinae<br />
Spiracular disc with five or fewer lobes . . . . . . . . . . . . . . . 2<br />
2 Body with prominent processes, at least half as long as <strong>the</strong> body width and arranged in<br />
longitudinal rows (figs 17- 20) . . . . . . . . . . . . . Cylindro<strong>to</strong>minae<br />
Body without prominent processes; at most with tubercles which are much shorter than half<br />
<strong>the</strong> body width (figs 21 - 29) . . . . . . . . . . . . . . . Limoniinae<br />
Tipulinae. The larvae <strong>of</strong> this subfamily include <strong>the</strong> familiar 'lea<strong>the</strong>r jackets' and are<br />
very similar in appearance: usually cylindrical, rarely depressed; with a <strong>to</strong>ugh unpigmented<br />
cuticle, usually clo<strong>the</strong>d in short, usually dense, pubescence. The colour may<br />
vary with <strong>the</strong> type <strong>of</strong> soil in which <strong>the</strong> larvae are found. There are invariably 6 anal lobes<br />
around <strong>the</strong> posterior spiracles. These lobes vary in size and degree <strong>of</strong> sclerotization<br />
according <strong>to</strong> <strong>the</strong> habitat in which <strong>the</strong> larvae are found. Soil is <strong>the</strong> main larval habitat<br />
with divergence in<strong>to</strong> decaying wood (Ctenophora) and mosses (Dolichopeza) , while<br />
some are aquatic (Prionocera, Tipula maxima Poda). In pasture soil some species attack<br />
<strong>the</strong> roots <strong>of</strong> cereals, root crops, vegetables such as Brassica, and also damage many<br />
35
herbaceous garden plants. The main pest species is Tipula paludosa Meigen which<br />
occurs in all soil types (clay, marl, peat, sand). The worst attacks occur in <strong>the</strong> spring,<br />
especially on young plants <strong>of</strong> crops sown on freshly ploughed grassland when <strong>the</strong><br />
lea<strong>the</strong>r jacket population may be as high as hundreds <strong>of</strong> thousands per acre. Usually<br />
Jea<strong>the</strong>rjackets feed on roots and underground stems just below <strong>the</strong> soil surface but on<br />
warm nights <strong>the</strong>y will feed on <strong>the</strong> surface, cutting <strong>of</strong>f <strong>the</strong> plants at soil level. Sometimes,<br />
especially after heavy rain, <strong>the</strong>y will be forced up through cracks in concrete, between<br />
paving s<strong>to</strong>nes etc and become trapped on <strong>the</strong> surface in very large numbers on hard<br />
areas in<strong>to</strong> which <strong>the</strong>y cannot burrow, such as paths and concrete surrounds <strong>of</strong> houses,<br />
and thus be reported as a nuisance.<br />
Brindle (I 967) keys <strong>the</strong> genera and species <strong>of</strong> this subfamily. The common 'pest'<br />
species usually found in pasture soil or gardens may be identified as follows.<br />
Dorsum <strong>of</strong> prothorax with two raised welts (fig. 16). (N. appendiculata (Pierre) ( = maculata<br />
Meigen) is a common garden species) . . . . . Nephro<strong>to</strong>ma spp.<br />
Dorsum <strong>of</strong>prothorax without raised welts (fig. 15). . . . . (Tipula sp.) 2<br />
2 Dorsal pairs (dl) <strong>of</strong> anal lobes sclerotized <strong>to</strong> tip (fig. 13) . . . T. vernalis Meigen<br />
Dorsal lobes not sclerotized . . . . . . . . . . . . . . . . . 3<br />
3 Only <strong>the</strong> lateral pair (lp) <strong>of</strong> anal papillae elongated, <strong>the</strong> ventral (vp) being rounded (fig. <strong>10</strong>)<br />
. . . . . . . . . . . . . . . . . . . . . T. paludosa Meigen<br />
At least two pairs <strong>of</strong> elongated, tapering, anal papillae, lateral (lp) and ventral (vp) . 4<br />
4 Pigmented marks between spiracles obvious (fig. 12), pigmented borders <strong>of</strong> anal lobes<br />
darker, disc whitish yellow, larva yellowish brown in colour, not translucent . . . .<br />
. . . . . . . . . . . . . . . . . . . . T. czizeki De Jong<br />
Pigmented marks between spiracles weak, <strong>of</strong>ten absent (fig. 11), pigmented borders <strong>of</strong> anal<br />
lobes lighter, disc whitish, larva greyish, ra<strong>the</strong>r translucent . . . . T. oleracea L.<br />
Cylindro<strong>to</strong>minae. The four <strong>British</strong> species <strong>of</strong> this subfamily have very distinctive<br />
caterpillar-like larvae which feed openly on <strong>the</strong> leaves <strong>of</strong> marsh plants.<br />
Diogma glabra/a (Meigen) (fig. 17) is green in colour and occurs in terrestrial mosses,<br />
usually on s<strong>to</strong>nes in limes<strong>to</strong>ne woodlands; Triogma trisulcata (Schummel) is dark<br />
brown or brownish green and occurs in semi-aquatic mosses (e.g. Hypnum cuspidatum)<br />
on moors; Phalacrocera replicata (L.) (fig. 19) is dark brown, paler below and is found<br />
on aquatic mosses (e.g. Hypnumfluitans or Sphagnum) in acid pools on moors or in<br />
woodlands; Cylindro<strong>to</strong>ma distinctissima Meigen is light green, darker dorsally and<br />
feeds on <strong>the</strong> leaves <strong>of</strong> Caltha, Viola, Stellaria, <strong>An</strong>emone, etc.<br />
Brindle ( 1967) keys <strong>the</strong> <strong>British</strong> genera and species.<br />
Limoniinae. The larvae <strong>of</strong> this subfamily can be approximately separated in<strong>to</strong> four<br />
tribes as below.<br />
Brindle ( 1967) keys <strong>the</strong> <strong>British</strong> genera and species so far described as larvae and Crisp<br />
& Lloyd (1954) give much useful biological information.<br />
Limoniini - larvae without obvious anal lobes, frequently living in silken tubes,<br />
herbivorous. Larvae occur in wet earth, decaying vegetation immersed in water or<br />
mud, on partly submerged s<strong>to</strong>nes in shallow streams, on piles and bridge piers where<br />
<strong>the</strong>y feed on algae. Larvae <strong>of</strong> Limonia (Geranomyia) unicolor Haliday feed on algae on<br />
marine structures washed by <strong>the</strong> tide. L. nubeculosa Meigen (fig. 29) is a very common<br />
species beneath leaves in woods. There are 58 <strong>British</strong> species in 7 genera in this tribe <strong>of</strong><br />
which some 25 species <strong>of</strong> Limonia s. I. are unknown in <strong>the</strong> larval stage.<br />
Pediciini - larvae with two anal lobes, free living and carnivorous. The larva <strong>of</strong><br />
Pedicia rivosa L. (fig. 25) has four pairs <strong>of</strong> creeping welts or tubercles which lack ventral<br />
hook-like spines at <strong>the</strong> tip. It is fairly common is aquatic or semi-aquatic mosses or in<br />
decaying vegetation in boggy places near streams and springs, where it feeds mainly<br />
on Diptera larvae (e.g. Chironomidae, Cera<strong>to</strong>pogonidae). <strong>An</strong>o<strong>the</strong>r common species<br />
Dicranota bimaculata Schummel has 5 pairs <strong>of</strong> well developed ventral pseudopods,<br />
36
which have prominent curved rows <strong>of</strong> hook-like spines at <strong>the</strong> tip (Figs 22-23). It is<br />
found in aquatic mosses or in marshy soils, mud or gravel near streams and rivers,<br />
where it feeds mainly on small worms (Oiigochaeta). Uta occurs in fungi. There are 19<br />
<strong>British</strong> species in <strong>the</strong> 3 genera in this tribe <strong>of</strong> which some species <strong>of</strong> Dicranota are<br />
unknown in <strong>the</strong> larval stage.<br />
Hexa<strong>to</strong>mini -larvae with 4 anal lobes, free living and carnivorous. The larvae <strong>of</strong><br />
Hexa<strong>to</strong>ma (fig. 27) are greenish and occur in sand by <strong>the</strong> edges <strong>of</strong> rivers and streams and<br />
share with <strong>the</strong> subgenera Phylidorea and Elaeophila <strong>of</strong> Limnophila and some Pi/aria<br />
species <strong>the</strong> characteristic swollen penultimate abdominal segment (though this may<br />
occur only in preserved larvae). There are 42 <strong>British</strong> species in 8 genera <strong>of</strong> which <strong>the</strong><br />
larvae <strong>of</strong> 11 species (4 Paradelphomyia, 2 Pi/aria, 5 Limnophila s. /.)are unknown.<br />
Eriopterini - larvae with 5 anal lobes, free living, herbivorous. This, <strong>the</strong> largest<br />
tribe, is <strong>the</strong> least well known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> because <strong>of</strong> <strong>the</strong>ir smaller size<br />
and burrowing habits. However, <strong>the</strong>y can occasionally occur in large numbers, e.g.<br />
Rhypholophus haemorrhoidalis Zetterstedt (almost black in colour) and Erioptera lutea<br />
Meigen. The larvae and pupae <strong>of</strong> E. squalida Loew have piercing spiracles<br />
with which <strong>the</strong>y obtain oxygen from <strong>the</strong> roots <strong>of</strong> aquatic plants (Glyceria maxima)<br />
(Houlihan, 1969). Of <strong>the</strong> 96 <strong>British</strong> species in 14 genera only 18 species in 9 genera are<br />
known as larvae.<br />
Psychodidae<br />
(Figs: larvae 30-40, egg 974, pupa 1112)<br />
The family Psychodidae (moth-flies) contains some 1200 world species <strong>of</strong> which 84<br />
species representing some 16 genera occur in <strong>the</strong> <strong>British</strong> Isles. Larvae <strong>of</strong> <strong>the</strong> <strong>British</strong><br />
species may be grouped in<strong>to</strong> three sub-families and separated as follows.<br />
Abdomen ending in a tubular siphon which bears <strong>the</strong> posterior spiracles at its apex and a pair<br />
<strong>of</strong> fan-like brushes (figs 30-37) . . . . . . . . . . . . . . Psychodinae<br />
Abdomen not ending in a tubular siphon; posterior spiracles on each side <strong>of</strong> 8th abdominal<br />
segment . . . . . . . . . . . . . . 2<br />
2 Body without conspicuous setae (fig. 40) . . . . . . . . . . . Trichomyiinae<br />
Body with conspicuous setae (fig. 39) . . . . . . . . . . . Sycoracinae<br />
Psychodinae. The genus Psychoda contains 15 <strong>British</strong> species <strong>of</strong> which <strong>the</strong> larvae <strong>of</strong><br />
13 are described and keyed by Satchell (1947). Psychoda alternata Say, Ps. albipennis<br />
Zetterstedt) ( =severini Tonnoir) and Ps. cinerea Banks are frequently found in sewage<br />
bacteria beds. Ps. alternata (figs. 30, 31) is known as <strong>the</strong> trickling filter fly (see also<br />
under Phoridae) and <strong>the</strong> larvae also occur in rotting seaweed and <strong>the</strong> sodden refuse <strong>of</strong><br />
disused farm feeding troughs, washings from animal cages, kitchen sink U-traps, etc.; it<br />
appears <strong>to</strong> show a preference for foul and wet habitats but not cow dung. Several<br />
species <strong>of</strong> Psychoda regularly breed in cow dung and o<strong>the</strong>rs occur in decaying<br />
vegetation. The common Ps. albipennis (figs 32-33) has been reared from horse and<br />
cow dung and rotting carrots; it has also been involved in cases <strong>of</strong> human urogenital<br />
myiasis (Zumpt, 1965). The larva <strong>of</strong> Ps. brevicornis Tonnoir (figs 35- 36) is adapted <strong>to</strong><br />
feeding on <strong>the</strong> surface <strong>of</strong> dung and differs from o<strong>the</strong>r dung-feeding species in having a<br />
lateral fringe <strong>of</strong> hairs <strong>to</strong> keep it afloat on <strong>the</strong> semi-liquid medium and only a short<br />
respira<strong>to</strong>ry siphon. Ps. surcouji Tonnoir is ano<strong>the</strong>r cow dung species, but larvae occasionally<br />
occur in milk bottles (see also Phoridae and Drosophilidae for this habit).<br />
Ps. lobata Tonnoir occurs in fungi.<br />
Satchell ( 1949) described 15 <strong>of</strong> <strong>the</strong> 24 <strong>British</strong> species <strong>of</strong> Pericoma <strong>the</strong>n recognised but<br />
Vaillant (1971-1976) has since transferred several species <strong>to</strong> o<strong>the</strong>r genera. The larvae<br />
are heavily pigmented and occur amongst clumps <strong>of</strong> damp moss and algal mats bordering<br />
<strong>the</strong> margins <strong>of</strong> streams where several species may be found in numbers quite close<br />
37
<strong>to</strong>ge<strong>the</strong>r. P. trivia/is Ea <strong>to</strong>n (figs 37- 38) is a common species, <strong>the</strong> larvae <strong>of</strong> which may be<br />
seen creeping about on <strong>the</strong> surface <strong>of</strong> mud, though <strong>the</strong> mud particles trapped in <strong>the</strong><br />
dorsal setae may render <strong>the</strong>m inconspicuous apart from <strong>the</strong> winding tracks left by<br />
<strong>the</strong>ir movements. The larvae <strong>of</strong> P. trifasciata (Meigen) and P. calcilega Feuerborn are<br />
normally thickly encrusted with lime.<br />
The larva <strong>of</strong> Philosepedon humeralis (Meigen) (fig. 34), <strong>the</strong> only <strong>British</strong> representative<br />
<strong>of</strong> <strong>the</strong> genus, feeds in dead snails (Vaillant, 1961; Smith & Grensted, 1963).<br />
Only three <strong>of</strong> <strong>the</strong> 17 <strong>British</strong> species <strong>of</strong> Telma<strong>to</strong>scopus are known in <strong>the</strong> larval stage;<br />
<strong>the</strong>y should be sought at <strong>the</strong> edges <strong>of</strong> ponds or lakes where <strong>the</strong> ground is moist and<br />
occasionally submerged, though Wi<strong>the</strong>rs (1986) records T. tristis Meigen and T.<br />
advenus Ea<strong>to</strong>n from tree rot-holes.<br />
Jung (1956) keys and describes some Panimerus, Threticus and Cly<strong>to</strong>cerus larvae.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> o<strong>the</strong>r <strong>British</strong> genera and species remain undescribed from<br />
native material but Vaillant (1971-1976) should be consulted for descriptions <strong>of</strong> some<br />
and notes on <strong>the</strong> likely habitats in which <strong>to</strong> search for o<strong>the</strong>rs (but frequently under<br />
names differing from <strong>the</strong> <strong>British</strong> Check List).<br />
Trichomyiinae. The larva <strong>of</strong> Trichomyia urbica Curtis, <strong>the</strong> only <strong>British</strong> species, is<br />
found in <strong>the</strong> rotting wood <strong>of</strong> fallen trees where it makes a gallery running in <strong>the</strong><br />
direction <strong>of</strong> <strong>the</strong> grain (Keilin & Tate 1937). The body, unlike that <strong>of</strong> o<strong>the</strong>r Psychodidae,<br />
is devoid <strong>of</strong> long setae and <strong>the</strong> head is strongly chitinized (fig. 40).<br />
Sycoracinae. The larvae <strong>of</strong> this subfamily are associated with fast streams and waterfalls<br />
where <strong>the</strong>y live on rocks in <strong>the</strong> splash zone or near <strong>the</strong> surface. Sycorax silacea<br />
Curtis (fig. 39), <strong>the</strong> only <strong>British</strong> representative <strong>of</strong> <strong>the</strong> subfamily, resembles a small water<br />
louse (Asellus) and is found among mosses or rotting leaves in such situations.<br />
<strong>An</strong> account <strong>of</strong> <strong>the</strong> <strong>British</strong> species <strong>of</strong> Psychodidae, by Mr P. Wi<strong>the</strong>rs, is in press.<br />
Ptychopteridae<br />
(Figs: larva 41 , pupa 1113)<br />
The family Ptychopteridae contains more than 60 world species in three genera. In<br />
~ritain <strong>the</strong> family is represented by only <strong>the</strong> genus Ptychoptera with 7 species <strong>of</strong> which 6<br />
have been described in <strong>the</strong> larval stage.<br />
The larva (fig. 41) is very distinctive, with a long, narrow cylindrical body<br />
terminating in a long retractile respira<strong>to</strong>ry siphon. It is whitish-grey in colour,<br />
frequently with a pair <strong>of</strong> brownish eye-spots. The respira<strong>to</strong>ry system is metapneustic.<br />
Small prolegs, each with a single hook-like spine (vh) are present on abdominal segments<br />
1- 3. The presence <strong>of</strong> lateral and median brushes on <strong>the</strong> labrum, and <strong>of</strong> setal<br />
brushes on <strong>the</strong> mandibles (md) and maxillae suggest a detritus feeding habit.<br />
Ptychoptera larvae are semi-aquatic and inhabit saturated mud, or shallow muddy<br />
pools in marshes, Sphagnum pools, or <strong>the</strong> margins <strong>of</strong> streams. Larvae <strong>of</strong> P. contaminata<br />
(L.) prefer mud covered with water and having a high organic content; <strong>the</strong>y are usually<br />
common around farmyard pools and similar situations. P. albimana(F.)is usually found<br />
in mud without a high organic content, e.g. open marshes, thin woodland; P. lacustris<br />
Meigen and P. pa/udosa Meigen (and P. longicauda Tonnoir teste R.l. Vane-Wright)<br />
occur most frequently in woodland streams, in medium or dense shade; P. scutel/aris<br />
Meigen is restricted <strong>to</strong> acid pools with Sphagnum, <strong>of</strong>ten at high altitudes in upland<br />
marshes; P. minuta Tonnoir has been found in a drainage channel around peaty fields.<br />
Brindle (1962d, 1966) and Hansen (1981) provide keys <strong>to</strong> <strong>the</strong> known <strong>British</strong> larvae<br />
with descriptions and ecological notes and Miall ( 1985) gives some interesting<br />
behavioural notes.<br />
38
Dixidae<br />
(Figs: larva 42, pupa 1114)<br />
Most <strong>of</strong> <strong>the</strong> 200 known species <strong>of</strong> Dixidae are Holarctic. Three genera are<br />
recognised, two <strong>of</strong> which occur in Britain: Dixa with 6 species and Dixella with 8<br />
species. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> all <strong>British</strong> species are known.<br />
The larvae are aquatic and are found at <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water which has earned<br />
<strong>the</strong>m <strong>the</strong> common name <strong>of</strong> meniscus midges. The resting larva adopts a U-shaped<br />
position (fig. 42) and joins <strong>the</strong> drift when disturbed. They are usually in contact with a<br />
plant, leaf, s<strong>to</strong>ne or piece <strong>of</strong> wood, where <strong>the</strong>y feed on microscopic organic particles<br />
and microorganisms swept in by <strong>the</strong> labral brushes. Dixa is usually found in running<br />
water (though always marginal) and Dixella in still water, while both genera occur<br />
in slow-moving water. Dixa larvae sometimes form a significant component <strong>of</strong> <strong>the</strong><br />
invertebrate fauna occurring in <strong>the</strong> downstream drift in rivers and streams.<br />
Disney (I 975) keys <strong>the</strong> <strong>British</strong> species in <strong>the</strong> larval, pupal and adult <strong>stages</strong>. The two<br />
genera may be separated as follows.<br />
Usually five (3- 7) or six (2- 7) <strong>of</strong> <strong>the</strong> abdominal segments each with a dorsal ring or 'crown' <strong>of</strong><br />
hairs (fig. 42,r) (poorly developed along anterior and posterior sec<strong>to</strong>rs in D. puberula<br />
Loew) . . . . . . . . . Dixa<br />
Abdominal segments without dorsal rings <strong>of</strong> setae . . . . . . . . . . DixeUa<br />
Chaoboridae<br />
(Figs: larvae 45-46, eggs 975, pupa 1115)<br />
Most <strong>of</strong> <strong>the</strong> <strong>10</strong>0 or so known species <strong>of</strong>Chaoboridae occur in <strong>the</strong> Neotropical region.<br />
The group was formerly regarded as a subfamily <strong>of</strong> <strong>the</strong> Culicidae but is now almost<br />
universally accepted as separate family. Three subfamilies are recognised, only one<br />
<strong>of</strong> which (Chaoborinae) occurs in Britain, where it is represented by two genera,<br />
Chaoborus with 4 species and M ochlonyx with 2 species.<br />
Chaoborus larvae have long been known as 'phan<strong>to</strong>m' larvae because <strong>of</strong> <strong>the</strong><br />
remarkably transparent body. The larvae <strong>of</strong> Mochlonyx are less transparent than<br />
Chaoborus. Hydrostatic organs enable chaoborid larvae <strong>to</strong> float in a horizontal<br />
position in mid-water waiting for <strong>the</strong>ir prey (mosqui<strong>to</strong> and mayfly larvae, Daphnia,<br />
Cypris, etc). The larvae <strong>of</strong> Chaoborus are found in a wide variety <strong>of</strong> lakes and ponds<br />
while Mochlonyx larvae prefer pools in woodland (M. culiciformis De Geer) or acid<br />
moorland (M. fuliginosus Felt). Larvae <strong>of</strong> <strong>the</strong> two genera may be separated by <strong>the</strong><br />
following couplet.<br />
Thorax not greatly swollen, and not much wider than abdomen; head longer and narrower,<br />
eyes smaller; anal segment without siphon; respira<strong>to</strong>ry system apneustic (fig. 45) . . .<br />
. . . . . . . . . . . . . Chaoborus Lichtenstein<br />
Thorax greatly swollen and much wider than abdomen (fig. 46); head shorter and broader,<br />
eyes larger, anal segment with siphon; respira<strong>to</strong>ry system metapneustic . . .<br />
. . . . . . . . . . . . . . . . . . . . . . Mochlonyx Loew<br />
Brindle ( 1962) keys <strong>the</strong> larvae <strong>of</strong> all <strong>the</strong> <strong>British</strong> species <strong>of</strong> <strong>the</strong> family and gives notes<br />
on <strong>the</strong>ir ecology, and Miall ( 1895) also gives an account <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Culicidae<br />
(Figs: larvae 43, 44; eggs 976-978; pupa 1116; adult emergence 1329)<br />
Over 3000 species <strong>of</strong> true mosqui<strong>to</strong>es have been described and three subfamilies <strong>of</strong><br />
worldwide distribution are recognised. Two subfamilies occur in Britain: <strong>An</strong>ophelinae,<br />
39
epresented by a single genus (<strong>An</strong>opheles) with 5 species, and Culicinae, with 5 genera<br />
and 27 species. Larvae <strong>of</strong> <strong>the</strong> two subfamilies found in Britain may be distinguished as<br />
follows:<br />
Eighth abdominal segment without an elongate dorsal siphon (fig. 43), larvae resting<br />
horizontal <strong>to</strong> <strong>the</strong> water surface . . . . . . . . . . . . . . <strong>An</strong>ophelinae<br />
Eighth abdominal segment with a dorsal siphon which is longer than broad (fig. 44), larvae<br />
resting obliquely <strong>to</strong> <strong>the</strong> surface . . . . . . . . . . . . . . . Culicinae<br />
The final stage (fourth ins tar) larvae <strong>of</strong> <strong>the</strong> <strong>British</strong> species are keyed by Snow ( 1984);<br />
Crans<strong>to</strong>n et al. ( 1987) provide keys <strong>to</strong> fourth ins tar larvae, pupae and adults.<br />
All mosqui<strong>to</strong> larvae are aquatic but occur in a variety <strong>of</strong> habitats and ecologically<br />
may be referred <strong>to</strong> four groups.<br />
I. Domestic species (adults overwinter indoors): <strong>An</strong>opheles atroparvus van Thiel (brackish<br />
water), A. messeae Falleroni (permanent ponds, slow streams), Culiseta ( = Theobaldia)<br />
annulata Schrank (foul or brackish water in stagnant ponds, tanks. waterbutts, etc.),<br />
Culex pipiens L. typical form (natural and artificial collections <strong>of</strong> fresh, foul or brackish<br />
water), C. pipiens form molestus Forskal (usually in water below ground level in semi<br />
darkness).<br />
2. Salt-marsh species (on or near coasts): Aedes caspius Pallas (brackish water, also in sewage<br />
effluents), A . detritus (Haliday) (brackish or saline water), A. dorsalis (Meigen) (brackish<br />
water).<br />
3. Arboreal species (tree-holes): <strong>An</strong>opheles plumbeus Stephens, Aedes geniculatus (Oiivier),<br />
Orthopodomyia pulcripalpis (Rondani).<br />
4. Rural species (non-domestic, in ground waters usually away from coasts): all <strong>the</strong> remaining<br />
species.<br />
Some species on <strong>the</strong> <strong>British</strong> List have only been recorded once or twice, but because<br />
<strong>of</strong> <strong>the</strong>ir medical importance are worthy <strong>of</strong> fur<strong>the</strong>r attention. Aedes aegypti (L.) is <strong>the</strong><br />
principal vec<strong>to</strong>r <strong>of</strong> urban yellow fever abroad and although temperature requirements<br />
preclude it from becoming established in Britain under natural conditions, <strong>the</strong>re was a<br />
local outbreak <strong>of</strong> yellow fever in Swansea in 1865, almost certainly from a single<br />
generation introduced from a ship. The species is extensively cultured in labora<strong>to</strong>ries<br />
and escapes may occur. Larvae <strong>of</strong> <strong>An</strong>opheles algeriensis Theobald have been recorded<br />
only once from three localities in Norfolk where <strong>the</strong>y occurred (and persisted for some<br />
years) in extensive shallow puddles among thick sedges on <strong>the</strong> marshes adjoining<br />
Hickling Broad. This species is doubtfully indigenous but investigations by a resident<br />
in <strong>the</strong> area would be worthwhile as it is one <strong>of</strong> <strong>the</strong> species capable <strong>of</strong> transmitting<br />
malaria and <strong>the</strong> disease used <strong>to</strong> occur in East <strong>An</strong>glia. The species has recently been<br />
discovered in <strong>An</strong>glesey (Morgan, 1987). Locally contracted cases <strong>of</strong> malaria can still<br />
occur in <strong>the</strong> U.K. in <strong>the</strong> vicinity <strong>of</strong> airports when mosqui<strong>to</strong>es are inadvertantly<br />
imported in aircraft. Aedes leucomelas Meigen has only been recorded once from<br />
Britain (Widmerpool, Notts, May 1919) and Culex modestus Ficalbi has only been<br />
recorded in 1944-1945 in <strong>the</strong> Portsmouth area.<br />
Apart from <strong>the</strong>se 'exotic' species <strong>the</strong>re is still much <strong>to</strong> be learned about <strong>the</strong> larvae <strong>of</strong><br />
most mosqui<strong>to</strong>es, especially on <strong>the</strong>ir precise feeding habits and distribution.<br />
The larvae <strong>of</strong> <strong>An</strong>opheles collect <strong>the</strong>ir food at <strong>the</strong> surface and <strong>of</strong> Culicines below <strong>the</strong><br />
water surface. Two different methods <strong>of</strong> feeding are adopted: by nibbling algae, dead<br />
leaves, etc. (e.g. Aedes, except A. cinereus Meigen) or by movements <strong>of</strong> <strong>the</strong> mouth<br />
brushes (figs 43, 44, br) which create a whirlpool bringing floating organic particles in<strong>to</strong><br />
<strong>the</strong> mouth (remaining genera and species).<br />
Mattingly (1969) has drawn attention <strong>to</strong> <strong>the</strong> interesting problem <strong>of</strong> mouthbrush<br />
polymorphism and particle size <strong>of</strong> food taken, which can apparently vary during a<br />
life-his<strong>to</strong>ry and requires fur<strong>the</strong>r investigation.<br />
40
There is an active mosqui<strong>to</strong> recording scheme (organiser Dr Keith Snow, Nor<strong>the</strong>ast<br />
London Polytechnic) which prospective researchers on <strong>the</strong> family are recommended <strong>to</strong><br />
support.<br />
The eggs <strong>of</strong> <strong>An</strong>ophelinae have floats and are laid singly (fig. 978). Eggs <strong>of</strong>Culicinae<br />
lack floats and may be laid singly on soil etc. in seasonally dry pools (Aedes, fig 977) or<br />
in batches or rafts on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water (Culex, fig 976). Chris<strong>to</strong>phers ( 1945) has<br />
written on <strong>the</strong> structure and function <strong>of</strong> Culex egg rafts.<br />
The pupa (fig. 1116) is comma shaped and swims with violent lashing movements <strong>of</strong><br />
<strong>the</strong> abdomen. It rests at <strong>the</strong> surface where it takes in air through respira<strong>to</strong>ry trumpets<br />
which extend internally in<strong>to</strong> <strong>the</strong> thoracic spiracles. In <strong>An</strong>ophelinae <strong>the</strong> trumpets are<br />
more or less conical; in Culicinae <strong>the</strong>y are more or less cylindrical.<br />
While this Handbook was in press an important work on <strong>the</strong> larval habitats <strong>of</strong><br />
mosqui<strong>to</strong>es appeared (Laird, 1988).<br />
Thaumaleidae<br />
(Figs: larva 4 7, pupa 1117)<br />
The 60 known species <strong>of</strong> Thaumaleidae are mostly Holarctic in distribution. Five<br />
genera are recognised <strong>of</strong> which only one, Thaumalea, is represented in Britain by 3<br />
species.<br />
The <strong>immature</strong> <strong>stages</strong> are aquatic. The larvae (fig. 47) resemble those <strong>of</strong>Chironomidae<br />
but have spiracles on <strong>the</strong> prothoracic and terminal abdominal segments and <strong>the</strong><br />
thoracic prolegs are unpaired (see key). They are found where clean water flows in thin<br />
films over rocks so that <strong>the</strong>ir backs are exposed above <strong>the</strong> surface. They feed on<br />
vegetable debris and dia<strong>to</strong>ms. Saunders (1923) describes <strong>the</strong> larva <strong>of</strong> Th. testacea<br />
Macquart but <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> o<strong>the</strong>r two species are undescribed.<br />
Cera<strong>to</strong>pogonidae<br />
(Figs: larvae 48- 52, egg 979, pupae 1118- 1120)<br />
About 4000 species <strong>of</strong> Cera<strong>to</strong>pogonidae (biting midges) have been described<br />
throughout <strong>the</strong> world, and 154 species in 17 genera (several divided in<strong>to</strong> numerous<br />
subgenera) occur in Britain. Four subfamilies are recognised <strong>of</strong> which Lep<strong>to</strong>conopinae<br />
does not occur in Britain. The larvae lack functional spiracles and respiration is<br />
cuticular or via anal blood-gills.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> most genera are aquatic and are found in streams, ponds,<br />
lakes, salt-marshes or water in tree holes. Some genera have terrestrial <strong>immature</strong><br />
<strong>stages</strong> and o<strong>the</strong>rs are semi-aquatic in wet soil, rotting wet wood or o<strong>the</strong>r decomposing<br />
vegetable matter. The adults <strong>of</strong> some genera are <strong>of</strong> medical and veterinary importance<br />
due <strong>to</strong> <strong>the</strong>ir habit <strong>of</strong> sucking vertebrate blood (e.g. Culicoides in Britain) and <strong>the</strong>ir<br />
potential role in disease transmission. Because <strong>of</strong> this, <strong>the</strong> larvae <strong>of</strong> <strong>the</strong> family,<br />
especially <strong>the</strong> blood-sucking genera, have attracted a certain amount <strong>of</strong> research but<br />
much remains <strong>to</strong> be done. The subfamilies occurring in Britain may be identified in <strong>the</strong><br />
larval <strong>stages</strong> by <strong>the</strong> following key.<br />
Thoracic prolegs present; body with conspicuous hairs <strong>of</strong>ten arising from raised tubercles<br />
(figs 48-49), <strong>of</strong>ten terrestrial . . . . . . . . . . . . . . Forcipomyiinae<br />
Thoracic pro legs absent; body without conspicuous body hairs or tubercles 2<br />
2 Posterior pro leg present with spines and hook lets (fig. 50). Usually crawling Dasyheleinae<br />
Posterior proleg absent, but last segment with hairs; body very smooth, whitish or<br />
translucent (figs 51- 52). Usually fast swimming with snake-like motion Cera<strong>to</strong>pogoninae<br />
Forcipomyiinae. This subfamily contains only two <strong>British</strong> genera, i.e. Forcipomyia<br />
(fig. 48) and Atrichopogon (fig. 49), which are mostly terrestrial (abroad, <strong>the</strong> subgenus<br />
41
Trichohelea <strong>of</strong> Forcipomyia is truly aquatic). The larvae <strong>of</strong> Forcipomyia have bulbous<br />
tips <strong>to</strong> some hairs (fig. 48, detail) within which a hygroscopic substance collects moisture<br />
from <strong>the</strong> atmosphere. The water flowing down <strong>the</strong> pedicel <strong>of</strong> <strong>the</strong> hair keeps <strong>the</strong> skin<br />
moist and facilitates cuticular respiration. These beads <strong>of</strong> moisture give <strong>the</strong> larva and<br />
pupa a striking appearance when first exposed. The larvae are gregarious (20-<strong>10</strong>0 or<br />
more) in habit and pupate <strong>to</strong>ge<strong>the</strong>r (fig. 1120). They are found in dark cavities where<br />
<strong>the</strong> atmosphere approaches saturation and favours <strong>the</strong> development <strong>of</strong> moulds and<br />
o<strong>the</strong>r fungi on which <strong>the</strong> larvae feed. There are 4 instars before pupation. Habitats<br />
include under bark (F. bipunctata L., F. picea Winnertz, F. nigra Winnertz), in rotting<br />
tap-roots <strong>of</strong> <strong>An</strong>gelica etc. (F. radicicola Edwards), rotting fungi (F. ciliata Winnertz),<br />
drying horse and cow manure (F. brevipennis Macquart). F. braueri (Wasmann) and F.<br />
myrmecophila (Egger) occur in ants' nests. F. pulchrithorax Edwards lives in <strong>the</strong><br />
granular, solidifying sap in open wounds on elm, chestnut and ash trees. Saunders<br />
( 1924) gives detailed descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> several species.<br />
Larvae <strong>of</strong> Atrichopogon (fig. 49) have <strong>the</strong> body segments flattened in cross section.<br />
They are found in moist soil at <strong>the</strong> water's edge and among accumulations <strong>of</strong> floating<br />
plants.<br />
The pupae <strong>of</strong> Forcipomyia and Atrichopogon retain <strong>the</strong> larval skin over <strong>the</strong> apical<br />
half <strong>of</strong> <strong>the</strong> body (fig. 1120). Few Diptera do this (see introduc<strong>to</strong>ry section on pupae).<br />
Dasyheleinae. This subfamily has only one genus, Dasyhelea (fig. 50).<br />
Keilin ( 1921) reviews <strong>the</strong> larval habitats, including those <strong>of</strong> <strong>the</strong> following <strong>British</strong><br />
species: D. bensoni Edwards (diplosis Keiffer, among filamen<strong>to</strong>us algae in a brackish<br />
pool); D. dufouri (Laboulbene) (in thick sap in wounds on elm); D.fiavifrons (Guerin<br />
Meneville) (in sap <strong>of</strong> beech and horse-chestnut); D. versicolor (Winnertz) (in humus<br />
around roots <strong>of</strong> Spiraea ulmaria and <strong>An</strong>gelica, from scum on surface <strong>of</strong> aquarium<br />
and (as obscura Winnertz) from elm sap, hornbeam sap and a water filled hole in an<br />
oak tree). D. bilineata Goetghebuer has been reared from <strong>the</strong> axil-waters <strong>of</strong> teasels<br />
(Dipsacusfullonum L.) by Disney & Wirth (1982). I have also seen Dasyhelea larvae<br />
from water in an old frying-pan used as a bird bath. The identity <strong>of</strong> <strong>the</strong> species recorded<br />
from water-filled tree-holes by Kitching (1972) is uncertain.<br />
Cera<strong>to</strong>pogoninae. This subfamily is divided in<strong>to</strong> five tribes. The Culicoidini contains<br />
Culicoides, <strong>the</strong> only genus occurring in Britain with blood-sucking adult females<br />
(divided in<strong>to</strong> six subgenera). The larvae (fig. 52) are long, smooth, dull whitish or<br />
translucent, progress with a snake-like lashing motion and are mostly aquatic.<br />
Kettle & Lawson ( 1952) key <strong>the</strong> larvae and pupae <strong>of</strong> 28 <strong>of</strong> <strong>the</strong> <strong>British</strong> species <strong>of</strong><br />
Culicoides, distinguish some o<strong>the</strong>r genera <strong>of</strong> <strong>the</strong> subfamily, and classify <strong>the</strong> larval<br />
habitats as follows (some genera from o<strong>the</strong>r tribes are also included):<br />
I. Bogland with ground cover <strong>of</strong> <strong>the</strong> mosses Sphagnum and Polytrichum penetrated by ]uncus<br />
articulatus (s. 1.): Culicoides impunctatus Goetghebuer (dominant), C. clin<strong>to</strong>ni Boorrnan<br />
(=C. truncorum Edwards, <strong>of</strong> Kettle & Lawson, 1952), C. heliophilus Edwards, C. albicans<br />
(Winnertz), C. obsoletus (Meigen) and Cera<strong>to</strong>pogon spp.<br />
2. Freshwater marsh without Sphagnum and Polytrichum, but with meadow plants e.g.<br />
Cardamine pratensis L. and Ranunculus spp. and sometimes with abundant ]uncus spp.:<br />
Culicoides kibunensis Tokunaga (=cubit a/is Edwards) (dominant), C. pallidicornis<br />
Kieffer, C. obsoletus, C. pulicaris (L.), C. delta Edwards, C. puncta/us (Meigen), C. odibilis<br />
A us ten, Stilobezzia spp., Cera<strong>to</strong>pogon spp. and Serromyia spp.<br />
3. Areas continually under water, e.g. swamp: C. kibunensis (dominant when Carex riparia<br />
cover present), C. odibilis (dominant when Carex cover absent), C. pulicaris. C. puncta/us.<br />
C.fascipennis (Staeger) (dominant in two swamps on margins <strong>of</strong> lochs).<br />
4. On areas <strong>of</strong> bare mud: C. odibilis (dominant), C. kibunensis, C. pallidicornis, C. pu/icaris. C.<br />
stigma (Meigen) and if <strong>the</strong>re is slight organic pollution C. nebeculosus (Meigen) is<br />
dominant.<br />
42
5. Coastal salt marshes: C. circumscriptus Kieffer and C. salinarius Kieffer (dominant on<br />
muddy sites), C. maritimus Kieffer and C. halophilus Kieffer (on vegetated sites).<br />
6. Dry cakes <strong>of</strong> cow dung: C. chiopterus (Meigen), C. dewulfi Goetghebuer ( = pseudochiopterus<br />
Downes & Kettle).<br />
Additional categories not included by Kettle & Lawson are as follows:<br />
7. Tree-holes: C.fagineus Edwards, C. truncorum Edwards, C. obsoletus (damp debris).<br />
8. Temporary pools in pasture: C. vexans (Staeger), C. brunnicans Edwards (small streams).<br />
9. Fungi (large Agaricaceae, Polyporaceae, Boletaceae): C. scoticus Downes & Kettle; C.<br />
obsoletus.<br />
<strong>10</strong>. Sap in tree wounds: C. chiopterus (on elm) (see also 6 above).<br />
The larvae <strong>of</strong> <strong>the</strong> remaining tribes should be sought as follows:<br />
Cera<strong>to</strong>pogonini: Cera<strong>to</strong>pogon and lsohelea in swampy banks <strong>of</strong> lakes, rivers and streams and in<br />
Sphagnum bogs.<br />
Stilobezziini: Monohelea in Sphagnum bogs and swampy parts <strong>of</strong> forest streams; Serromyia in<br />
margins <strong>of</strong> shallow waters and swamps;Stilobezzia in moist soil, <strong>of</strong>ten overgrown with moss<br />
along <strong>the</strong> margins <strong>of</strong> rivers and streams, silted up ditches, etc.<br />
Heteromyiini: C/inohelea in heavily vegetated overgrown meadows; sedgy boggy parts <strong>of</strong> flowing<br />
water in river flood plains.<br />
Sphaeromyiini: Mallochohelea inermis (Kieffer) and M. munda (Loew) in lakes and M. setigera<br />
(Loew) in rivers; Probezzia in rivers in sandy areas and among dense weeds; Sphaeromias in<br />
lakes (S.fasciatus Meigen in <strong>the</strong> depths).<br />
Palpomyiini: Palpomyia in lakes and rivers; Bezzia mostly in lakes, some in swamps and forest<br />
pools.<br />
There is plenty <strong>of</strong> scope for careful research on larval ecology and systematics in this<br />
important family. The biting adults <strong>of</strong> Culicoides can make life miserable for residents<br />
and visi<strong>to</strong>rs in nor<strong>the</strong>rn temperature regions (e.g. C. impunctatus in <strong>the</strong> highlands <strong>of</strong><br />
Scotland). A useful modern introduc<strong>to</strong>ry chapter on <strong>the</strong> family is provided by Kettle<br />
( 1984: 137- 158) and Atchley et al. ( 1981) give a full bibliography <strong>of</strong> Cera<strong>to</strong>pogonidae.<br />
Keys <strong>to</strong> genera are provided by Glukhova (1977, 1979).<br />
Chironomidae<br />
(Figs: larvae 53-67, eggs 980--982, pupa 1121)<br />
Over 5000 species <strong>of</strong> Chironomidae (non-biting midges) have so far been described<br />
throughout <strong>the</strong> world. Seven subfamilies are recognised, six <strong>of</strong> which are represented in<br />
Britain by over 450 species in 120 genera.<br />
Larval Chironomidae are <strong>the</strong> most abundant Diptera in aquatic ecosystems and<br />
<strong>of</strong>ten dominate <strong>the</strong> invertebrate fauna. Their occurrence in a wide range <strong>of</strong> ecological<br />
conditions has led <strong>to</strong> <strong>the</strong>ir use as indica<strong>to</strong>r organisms in classifying lakes, river<br />
zonation, water quality, etc. This has led <strong>to</strong> an intense study <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
Chironomidae in recent years. Chironomid larvae are a major source <strong>of</strong> food for<br />
freshwater fish and may constitute as much as 80% <strong>of</strong> <strong>the</strong>ir <strong>to</strong>tal food requirements.<br />
The larvae <strong>of</strong> some species (e.g. Metriocnemus hygropetricus Kieffer) play an essential<br />
role, along with Psychodidae, as browsers in trickling filter sewage beds. Occasionally<br />
larvae may occur in <strong>the</strong> domestic fresh water supply and par<strong>the</strong>nogenetic Paratanytarsus<br />
( = Stylotanytarsus), including apparently paedogenetic pupal forms, can complete<br />
<strong>the</strong>ir life cycles entirely within <strong>the</strong> system. (See introduc<strong>to</strong>ry sections on larvae, and<br />
agricultural and medical importance.)<br />
Not all Chironomidae have aquatic life-his<strong>to</strong>ries and <strong>the</strong> larvae <strong>of</strong> terrestrial species<br />
are not so well known. There is much scope for research in <strong>the</strong> elucidation <strong>of</strong> lifehis<strong>to</strong>ries<br />
providing that larvae are reared through <strong>to</strong> adults, males <strong>of</strong> which can now be<br />
43
eliably identified by <strong>the</strong> key in Pinder (1978). A typical pupa is illustrated (fig. 1121)<br />
and a key <strong>to</strong> pupal exuviae is provided by Lang<strong>to</strong>n (1984).<br />
Eggs are laid in a protective gelatinous matrix and each subfamily appears <strong>to</strong> have<br />
a characteristic shape <strong>of</strong> egg and arrangement within <strong>the</strong> mass, e.g. linear type<br />
(Orthocladiinae, Diamesinae); compact type- spherical, tear or fig-shaped, or<br />
occasionally attached <strong>to</strong> a solid object by a gelatinous stalk (Tanypodinae,<br />
Chironominae) (figs 980-982).<br />
The life-his<strong>to</strong>ries <strong>of</strong>Chironomidae are reviewed by Oliver (1971) and a bibliography<br />
<strong>of</strong> Chironomidae is provided by Fittkau et al. (1976). Keys <strong>to</strong> larvae and pupae are<br />
provided by various authors in Wiederholm ( 1983, 1986). These are cited below, under<br />
subfamilies. This comprehensive work will also eventually cover pupae and adults, and<br />
reliable keys beyond genus are cited.<br />
The subfamilies may be distinguished in <strong>the</strong> larval stage as follows (after Crans<strong>to</strong>n<br />
1982).<br />
<strong>An</strong>tenna retractile in<strong>to</strong> head capsule (fig, 53, an); ligula strongly developed (fig. 53.lg)<br />
. . . . . . . . . . . . . . . . . . . . . . . Tanypodinae<br />
<strong>An</strong>tenna not retractile; ligula not strongly developed . . . . . . . . . . . 2<br />
2 Premandibles absent; procerci 8- <strong>10</strong> times as high as wide (fig. 59, pc) . . Polloaominae<br />
Premandibles present (fig. 54, pm); procerci rarely more than 4 times as high as wide 3<br />
3 Third antennal segment annulate (fig. 56) . . . . . . . . Diarwsiww (Diamesini)<br />
Third antennal segment not annulate . . . . . . . . . . . . . . . . 4<br />
4 <strong>An</strong>terior part <strong>of</strong> labrum (fig. 54, la) with a transverse row <strong>of</strong>'scales' (fig. 57) . . . . .<br />
. . . . . . . . . . . . . . . . . . . . Diamesiaar (Protanypini)<br />
Labrum without this row <strong>of</strong> 'scales' . . . . . . . . . . . . . . . . 5<br />
5 Ventral part <strong>of</strong> <strong>the</strong> mentum (fig. 54, m) expanded laterally <strong>to</strong> form ventromental plates which<br />
are usually striated (fig. 55) . . . . . . . . . . . . . . Cllirooominae<br />
Ventromental plates not striated . . . . . . . . . . . . . . . . . 6<br />
6 Ventromental plates well developed, extending beyond mentum by at least half <strong>of</strong> <strong>the</strong> width<br />
<strong>of</strong> <strong>the</strong> flattened mentum . . . . . . . . . . . . . . . Prodiamesinae<br />
Ventromental plates smaller, never extending beyond mentum by as much as half width <strong>of</strong><br />
flattened mentum . . . . . . . . . . . . . . . . . . . . . . 7<br />
7 Hypopharynx (fig. 54, h) with an apical group <strong>of</strong> elongate, serrate hairs (fig. 58) . . . .<br />
. . . . . . . . . . . . . Tet.&<strong>to</strong>ge<strong>to</strong>ninae<br />
Hypopharynx with a few shorter hairs . . . . . . . . . . . . Ortllocladiinae<br />
Tanypodinae. Larvae <strong>of</strong> this subfamily are aquatic, may be red with haemoglobin,<br />
and do not construct tubes or cases. They prey on small Crustacea, oligochaete worms<br />
and <strong>of</strong>ten on <strong>the</strong> larvae <strong>of</strong> o<strong>the</strong>r Chironomidae. Adaptations <strong>to</strong> <strong>the</strong> predaceous habit<br />
include <strong>the</strong> retractile antennae and ra<strong>the</strong>r long stilt-like pro legs, <strong>the</strong> latter giving rise <strong>to</strong><br />
a rapid jerky motion (fig. 65). The pupae resemble those <strong>of</strong> mosqui<strong>to</strong>es and are freeliving<br />
active swimmers. Larvae <strong>of</strong> Tanypodinae are keyed (<strong>to</strong> genera) by Fittkau &<br />
Roback (in Wiederholm, 1983: 33- 1<strong>10</strong>).<br />
Podonominae and Diamesinae. Larvae <strong>of</strong> <strong>the</strong>se small subfamilies appear <strong>to</strong> be<br />
generally adapted for life in cold conditions and are found in mossy mountain springs<br />
and cold streams. Lasiodiamesa sphagnicola (Kieffer), however, favours wanner pools<br />
in Sphagnum bogs. Larvae <strong>of</strong> Podonominae feed on dia<strong>to</strong>ms, are free-living and very<br />
strong swimmers (even as pupae). Larvae <strong>of</strong> Podonominae are keyed by Brundin (in<br />
Wiederholm, 1983: 25- 31) and Diamesinae (<strong>to</strong> genera) by Oliver (in Wiederholm, 1983:<br />
115-138).<br />
Orthocladiinae. Some larvae <strong>of</strong> this subfamily are terrestrial, usually in moss covered<br />
soil. Larvae <strong>of</strong> Bryophaenocladius have been reported as attacking <strong>the</strong> roots <strong>of</strong> greenhouse<br />
plants, s<strong>to</strong>red pota<strong>to</strong>es, mosses, liverworts and commercial lettuce crops and are<br />
able <strong>to</strong> 'skip' like a cheese-skipper (Piophila) . I have seen larvae (probably Smittia sp.)<br />
44
eported as attacking: young plants <strong>of</strong> barley in Aberdeenshire (M.W. Shaw, pers.<br />
comm.); central shoots <strong>of</strong> winter corn in Lincolnshire (G. Murdoch, pers. comm.);<br />
winter wheat in Newcastle upon Tyne (4% <strong>of</strong> seed failed <strong>to</strong> germinate following 2 year<br />
ley, J.P. Rogerson, pers. comm.); and associated with rose-roots in large numbers in<br />
Reading.<br />
Many larvae <strong>of</strong> this subfamily have adapted <strong>to</strong> life in swift flowing rivers and<br />
streams (e.g. Rheocrico<strong>to</strong>pus, Eukie.fferiella). Cardiocladius larvae are free-living in<br />
waterfalls and swift currents where <strong>the</strong>y pupate in strong cocoons spun on rocks, s<strong>to</strong>nes<br />
and water-plants. Metriocnemus martinii Thienemann larvae are found in water filled<br />
tree holes (especially beech); M. ursinus Holmgren occurs in moss. Larvae <strong>of</strong> Clunio<br />
marinus Haliday, Halocladius spp. and Thalassosmittia thalassophila (Goetghebuer)<br />
are marine. Epoicocladius.flavens (Malloch) (=ephemerae Kieffer) larvae are found on<br />
mayfly nymphs which <strong>the</strong>y do not harm; <strong>the</strong> fully grown larva spins a gelatinous<br />
cocoon on <strong>the</strong> thorax <strong>of</strong> <strong>the</strong> mayfly nymph in which it pupates. Keys <strong>to</strong> <strong>the</strong> known<br />
larvae <strong>of</strong> Orthocladiinae are provided by Crans<strong>to</strong>n (1982) and Crans<strong>to</strong>n, Oliver &<br />
Sae<strong>the</strong>r (in Wiederholm, 1983: 149-291 ).<br />
Chironominae. Most larvae <strong>of</strong> this subfamily are red in colour, due <strong>to</strong> <strong>the</strong> presence <strong>of</strong><br />
haemoglobin which assists survival in low oxygen concentrations. They live in tubes <strong>of</strong><br />
mud or vegetable debris. In <strong>the</strong> tribe Chironomini <strong>the</strong> larvae are generally larger and<br />
include detritus feeders, filter feeders, symbionts, parasites and carnivores. The larvae<br />
<strong>of</strong> <strong>the</strong> common Chironomus plumosus L. and C. dorsalis Meigen live in mud in small<br />
stagnant pools. They construct fixed tubes <strong>of</strong> mud and vegetable debris respectively,<br />
but C. plumosus spins a sheet <strong>of</strong> salivary secretion across <strong>the</strong> tube and by undulations <strong>of</strong><br />
<strong>the</strong> body draws phy<strong>to</strong>plank<strong>to</strong>n and o<strong>the</strong>r particles in<strong>to</strong> <strong>the</strong> net (fig. 61) (filter feeder). C.<br />
dorsalis (fig. 66) and o<strong>the</strong>r Chironomus species only take food from <strong>the</strong> end <strong>of</strong> <strong>the</strong>ir<br />
tubes (detritus feeders). The larva <strong>of</strong> Glyp<strong>to</strong>tendipes paripes (Edwards) is found in <strong>the</strong><br />
mollusc <strong>An</strong>odonta (Beedham, 1966).<br />
Larvae <strong>of</strong> <strong>the</strong> tribe Tanytarsini are smaller in size and are detritus or filter feeders.<br />
Rheotanytarsus species are filter feeders that construct tubes <strong>of</strong> fine silt particles<br />
attached <strong>to</strong> s<strong>to</strong>nes and plants. They spin nets across arms at <strong>the</strong> entrance <strong>of</strong> <strong>the</strong> tube and<br />
<strong>the</strong> natural water current brings food particles in<strong>to</strong> <strong>the</strong> net, <strong>the</strong> water passing out<br />
through a hole at <strong>the</strong> end <strong>of</strong> <strong>the</strong> tube (figs 63- 64).<br />
Some larvae in <strong>the</strong> subfamily construct movable cases and adopt a mode <strong>of</strong> life<br />
similar <strong>to</strong> caddis larvae (e.g. Zavrelia, Stempellina and Lauterborniella). Larvae <strong>of</strong><br />
Chironominae are keyed <strong>to</strong> genus by Pindar & Reiss (in Weiderholm, 1983: 293-435).<br />
Telma<strong>to</strong>ge<strong>to</strong>ninae. Larvae <strong>of</strong> <strong>the</strong> two <strong>British</strong> species (Thalassomya frauenfeldii<br />
Schiner, Psamathiomya pectinata Deby) <strong>of</strong> this subfamily are adapted <strong>to</strong> marine<br />
conditions and live on intertidal rocks where <strong>the</strong>y feed on algae which <strong>the</strong>y scrape from<br />
<strong>the</strong> rocks. Females <strong>of</strong> Telma<strong>to</strong>ge<strong>to</strong>ninae are unique in <strong>the</strong> Chironomidae in laying<br />
eggs individually. Larvae <strong>of</strong> Telma<strong>to</strong>ge<strong>to</strong>ninae are keyed <strong>to</strong> genus by Crans<strong>to</strong>n (in<br />
Wiederholm, 1983: 17-22).<br />
Marine habitats have also been invaded by larvae <strong>of</strong>Orthocladiinae (e.g. Clunio) and<br />
Chironominae (e.g. Chironomus).<br />
Simuliidae<br />
(Figs: larvae 68--69, egg 985, pupa 1122)<br />
The family Simuliidae (black-flies) contains some 1450 species and has an almost<br />
cosmopolitan distribution. Two subfamilies are usually recognised, one <strong>of</strong> which<br />
(Simuliinae) is represented in Britain by 33 species contained in 3 genera (Simulium,<br />
Prosimulium and Metacnephia), but many sibling species, recognisable only on <strong>the</strong>ir<br />
45
chromosomes, are being found. The larvae are aquatic and usually found in running<br />
water including rapids and cascades. The three species <strong>of</strong> Prosimulium are found in hill<br />
streams. The commonest species <strong>of</strong> Simulium is S. ornatum Meigen, <strong>the</strong> larvae <strong>of</strong> which<br />
are found in all types <strong>of</strong>lowland water courses and are <strong>of</strong>ten abundant in small streams<br />
and flowing ditches. Simulium vernum Macquart and S. cryophilum Knoz are <strong>of</strong>ten<br />
abundant in many types <strong>of</strong> small streams. S. equinum (L.) is common, usually on rooted<br />
vegetation in large streams with a steady current and is frequently accompanied by S.<br />
lineatum (Meigen). S. reptans (L.) and S. tuberosum Lundstrom are common in larger<br />
s<strong>to</strong>ny rivers but <strong>the</strong> latter occurs usually above <strong>the</strong> limit <strong>of</strong> cultivation <strong>of</strong> <strong>the</strong><br />
surrounding land and <strong>the</strong> former below, though <strong>the</strong>re is an area <strong>of</strong> overlap. O<strong>the</strong>r<br />
species are more specialised in <strong>the</strong>ir habitat and consequently less common, e.g. S.<br />
costatum Friederichs (spring-fed streams near source), S. noelleri Friederichs (lake or<br />
pond outlets).<br />
The larvae are filter feeders on vegetable debris and dia<strong>to</strong>ms which <strong>the</strong>y strain from<br />
<strong>the</strong> water with <strong>the</strong>ir cephalic fans (figs 68-69). Occasionally <strong>the</strong>y may ingest small<br />
larvae such as Chironomidae or even early ins tars <strong>of</strong> <strong>the</strong>ir own kind. There are six <strong>to</strong><br />
eight larval instars in our species. There is a short pharate (see Introduction) stage<br />
(recognised by <strong>the</strong> much darker gill-spot) when <strong>the</strong> prepupa, still inside <strong>the</strong> last larval<br />
skin, spins its cocoon <strong>of</strong> silk, attached <strong>to</strong> a suitable substrate, inside which it pupates.<br />
The pupa has long filamen<strong>to</strong>us gills (fig. 1122).<br />
The eggs are eccentric ovoid in shape (fig. 985) and may be laid singly and loosely or<br />
in batches stuck <strong>to</strong> <strong>the</strong> substrate; <strong>the</strong>y are white when laid but darker later. Many<br />
species pass <strong>the</strong> winter as eggs but some do so as slowly growing larvae.<br />
Most adult female Simuliidae suck blood and <strong>the</strong>ir biting <strong>of</strong> man and animals causes<br />
irritation sometimes followed by <strong>to</strong>xic and allergic reactions. In tropical Africa and<br />
tropical America <strong>the</strong>y transmit <strong>the</strong> disease onchocerciasis (river blindness). In Britain<br />
<strong>the</strong>y are known <strong>to</strong> transmit <strong>the</strong> nema<strong>to</strong>de Onchocerca gutturosa <strong>to</strong> cattle (without<br />
serious results) and certain Pro<strong>to</strong>zoa (e.g. Leucocy<strong>to</strong>zoon) among birds. This medical<br />
and veterinary importance has led <strong>to</strong> a considerable amount <strong>of</strong> work on <strong>the</strong>ir <strong>immature</strong><br />
<strong>stages</strong> for <strong>the</strong> purpose <strong>of</strong> survey and control work throughout <strong>the</strong> world. The Simulium<br />
fauna <strong>of</strong> an area is best established by searching for <strong>the</strong>ir larvae and pupae in freshwater<br />
systems from small trickles <strong>to</strong> large rivers. Thorough surveys such as that <strong>of</strong>Crosskey<br />
(1985) in <strong>the</strong> London area would be worthwhile projects and add much <strong>to</strong> our<br />
knowledge <strong>of</strong> <strong>the</strong> ecology and distribution <strong>of</strong> <strong>the</strong> family.<br />
L. Davies ( 1968) keys <strong>the</strong> <strong>British</strong> species in <strong>the</strong> larval, pupal and adult <strong>stages</strong>, but <strong>the</strong><br />
keys <strong>to</strong> early <strong>stages</strong> are difficult <strong>to</strong> use and are currently under revision by <strong>the</strong><br />
Freshwater Biological Association. Ladle & Bass (1975) have described <strong>the</strong> larva <strong>of</strong> a<br />
new Metacnephia. The Diptera Check List in <strong>the</strong> Handbook series should be consulted<br />
for nomencla<strong>to</strong>rial changes.<br />
<strong>An</strong>isopodidae<br />
(Figs: larvae 70-73, egg 986, pupa 1123)<br />
The family <strong>An</strong>isopodidae contains about <strong>10</strong>0 species in six genera <strong>of</strong> which 5 species<br />
in 2 genera occur in Britain. Two subfamilies are recognised, <strong>the</strong> larvae <strong>of</strong> which may be<br />
separated as follows.<br />
<strong>An</strong>al segment ra<strong>the</strong>r truncate posteriorly; posterior spiracular disc larger and with distinct<br />
lobes (fig. 72); lines drawn through <strong>the</strong> axes <strong>of</strong> <strong>the</strong> diagonal posterior spiracles would<br />
intersect on disc . . . . . . . . . . . . . . . . . . <strong>An</strong>isopodinae<br />
One genus, Sylvicola ( = <strong>An</strong>isopus)<br />
<strong>An</strong>al segment elongated and more rounded posteriorly; posterior spiracular disc small,<br />
without distinct lobes; lines drawn through axes <strong>of</strong> posterior spiracles would not intersect<br />
on disc (fig. 73) . . . . . . . . . . . . . . . . Myce<strong>to</strong>biinae<br />
One genus and species, Myce<strong>to</strong>bia pallipes Meigen<br />
46
<strong>An</strong>isopodid larvae all have a perianal shield (<strong>of</strong> thickened hypodermal cells) on <strong>the</strong><br />
ventral surface <strong>of</strong> <strong>the</strong> anal segment, which spreads laterally with a sinuate margin (fig.<br />
71, S).<br />
The larvae <strong>of</strong> Sylvicola species (fig. 70) are yellowish in colour but <strong>the</strong> underlying<br />
organs impart a reddish tinge with dark brownish thoracic markings. They are usually<br />
associated with decaying vegetable matter such as pota<strong>to</strong>es, beet, decaying roots <strong>of</strong><br />
<strong>An</strong>gelica and burdock, mouldy decaying honeycombs, cardboard, fungi, decaying<br />
leaves in tree rot-holes, slime flux from tree wounds, but have also been found in cider<br />
and home-made wines, and liver that had been preserved in formalin for seven years<br />
(partly projecting and mouldy). Sylvicolafenestralis (Scopoli) has occurred in cases <strong>of</strong><br />
urogenital myiasis (Smith & Taylor, 1966, Morris, R.F. 1986). Sylvicola punctatus (F.)<br />
frequently occurs in dung. The remaining two <strong>British</strong> species are unknown in <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong>. The larvae <strong>of</strong> Sylvicola are <strong>of</strong>ten found in company with Sca<strong>to</strong>pse<br />
notata (L.) (Sca<strong>to</strong>psidae), Lonchaea chorea (F.) (Lonchaeidae), Myce<strong>to</strong>philidae and<br />
Sphaeroceridae.<br />
Larvae <strong>of</strong> M yce<strong>to</strong>bia pallipes Meigen (fig. 71) occur in slime or exudations on <strong>the</strong><br />
surface or under wet bark on decaying trees.<br />
The eggs <strong>of</strong> Sylvicola (fig. 986) and Myce<strong>to</strong>bia are enclosed in a gelatinous mass. The<br />
pupae are free and found in <strong>the</strong> same media as <strong>the</strong> larvae.<br />
Keilin & Tate ( 1940) give detailed descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Sylvicola<br />
fenestra/is and Myce<strong>to</strong>bia pallipes; Brindle ( 1962!) gives keys <strong>to</strong> species.<br />
Bibionidae<br />
(Figs: larvae 74-78, eggs 983- 984, pupa 1124)<br />
The family Bibionidae contains nearly 700 species in 8 genera throughout <strong>the</strong> world.<br />
Three subfamilies are recognized but only one <strong>of</strong> <strong>the</strong>se (Bibioninae) is present in<br />
Britain, represented by <strong>the</strong> two genera Bibio and Dilophus. Larvae <strong>of</strong> Bibio have two<br />
openings <strong>to</strong> <strong>the</strong> posterior spiracles and larger body processes in transverse rows (figs 74,<br />
78), while those <strong>of</strong> Dilophus have three spiracular openings (fig. 77) and shorter body<br />
processes except for one dorso-lateral series.<br />
Both genera are herbivorous, <strong>of</strong>ten on plant roots, and live in soil, wood debris,<br />
dung, etc. The larvae are slow moving and gregarious, being usually found clustered<br />
<strong>to</strong>ge<strong>the</strong>r.<br />
The eggs <strong>of</strong>Bibionidae (fig. 983) are laid in masses and appear ra<strong>the</strong>r opalescent. The<br />
newly hatched larva <strong>of</strong> Bibio (fig. 76) has conspicuous long hairs but that <strong>of</strong> Dilophus is<br />
short haired (fig. 75). There are four larval instars. Male pupae <strong>of</strong> Dilophus have 3<br />
pointed processes on <strong>the</strong> head and female pupae have a single process, but this is better<br />
developed than <strong>the</strong> single process present in <strong>the</strong> pupae <strong>of</strong> both sexes <strong>of</strong> Bibio (fig. 1124).<br />
Bibio larvae are usually found in woodland soil but B.johannis (L.) occurs in soil or<br />
dung in open country. Larvae <strong>of</strong> both genera are occasionally pests <strong>of</strong> cereal and root<br />
crops (see Economic section and Freeman & Lane (1985). Morris, H.M. (1922)<br />
suggested that such cases were due <strong>to</strong> larvae being carried in manure applied <strong>to</strong> <strong>the</strong> plants.<br />
Morris, H.M. (1917, 1921-1922) gives detailed descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Brindle (1962a) gives specific keys <strong>to</strong> <strong>the</strong> larvae so far known (2 <strong>of</strong> <strong>the</strong> 4 species <strong>of</strong><br />
Dilophus and 8 <strong>of</strong> <strong>the</strong> 14 <strong>of</strong> Bibio).<br />
Myce<strong>to</strong>philidae<br />
(Figs: larvae 79-86, 89-91; egg 988; pupa 1125)<br />
The family Myce<strong>to</strong>philidae (fungus gnats) contains about 3400 described species<br />
with an almost cosmopolitan distribution. In Britain about 450 species in 65 genera and<br />
all seven accepted subfamilies have been recorded.<br />
47
Larvae <strong>of</strong> six <strong>of</strong> <strong>the</strong> seven subfamilies (Manotinae not yet described) may be<br />
separated as follows (after Madwar 1937).<br />
Larva with one pair <strong>of</strong> thoracic and 8 pairs <strong>of</strong> abdominal spiracles (8th pair with horned<br />
prominences in Di<strong>to</strong>myia) (figs 79-80) . . . . . . . . . . . . Di<strong>to</strong>myiinae<br />
Larva with one pair <strong>of</strong> thoracic and at most 7 pairs <strong>of</strong> abdominal spiracles, or without<br />
spiracles (Keroplatinae) . . . . . . . . . . . . 2<br />
2 <strong>An</strong>tennae well developed, composed <strong>of</strong> three segments (fig. 81). Boli<strong>to</strong>philinae<br />
<strong>An</strong>tennae rudimentary, composed <strong>of</strong> one segment . 3<br />
3 Larva free; locomo<strong>to</strong>ry pads present (fig. 82) Myce<strong>to</strong>philinae<br />
Larva enclosed in slimy tube, or in web; pads absent 4<br />
4 Larva without spiracles, vermiform (worm-shaped) . Keroplatinae<br />
Larva with spiracles, not vermiform . . . . 5<br />
5 Larva peripneustic (fig. I) . . . . . . . . . most Sciophilinae<br />
Larva propneustic (fig. I) . . . . . . . . . . . . . . 6<br />
6 Epicranial plates meeting as usual on ventral surface (fig. 86) . . . . . . .<br />
. . . . . . . . . . . . . . . . (Genus Speolepta) Sciophilinae<br />
Epicranial plates not meeting on ventral surface . . . . Diadocidiinae<br />
The larvae <strong>of</strong>Myce<strong>to</strong>philidae are mostly found in <strong>the</strong> larger fungi and it is probable<br />
that even when found elsewhere <strong>the</strong> majority are feeding on micr<strong>of</strong>ungi or fungal<br />
hyphae.<br />
Chandler (in Stubbs & Chandler, 1978) lists fungus associations for 130 <strong>British</strong><br />
species <strong>of</strong> Myce<strong>to</strong>philidae and Hutson et al. (1980) give considerable information on<br />
life his<strong>to</strong>ries. Mad war (1937) and Las<strong>to</strong>vka ( 1971) describe some <strong>immature</strong> <strong>stages</strong>.<br />
There are three recognisable <strong>stages</strong> in <strong>the</strong> development <strong>of</strong> a s<strong>of</strong>t fungal fruiting body<br />
(sporophora): I, fresh with no signs <strong>of</strong> decay; 2, decaying through age and damage<br />
(mostly by fungivores); 3, semi-liquid, advanced state <strong>of</strong> decay (some <strong>to</strong>ugher fungi<br />
desiccate ra<strong>the</strong>r than rot and are unsuitable for fungus-gnats).<br />
Most myce<strong>to</strong>philids attack <strong>the</strong> first two <strong>stages</strong> but some species will continue <strong>to</strong><br />
develop in <strong>the</strong> last stage <strong>of</strong> almost any rotting fungus (e.g. Myce<strong>to</strong>philafungorum De<br />
Geer). O<strong>the</strong>r habits and habitats are discussed below under subfamilies.<br />
There is plenty <strong>of</strong> scope for research on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> provided care is taken<br />
<strong>to</strong> identify <strong>the</strong> fungi and <strong>the</strong> adult flies correctly. Adults <strong>of</strong> all subfamilies except<br />
Myce<strong>to</strong>philinae are keyed in Hus<strong>to</strong>n et al. (1980); o<strong>the</strong>r taxonomic literature is cited in<br />
that work.<br />
Di<strong>to</strong>myiinae. There are only two genera <strong>of</strong> this subfamily in Britain, each with a<br />
single species. The characteristic larvae <strong>of</strong> Di<strong>to</strong>myia fasciata Meigen (fig. 80) have<br />
been reared from a variety <strong>of</strong> hard polyporaceous fungi including Coriolus,<br />
Fomes, Daedalea, Polyporus, Bjerkandera, Trametes, lnonotus, Lep<strong>to</strong>porus. Symmerus<br />
annulatus (Meigen) (fig. 79) is found in rotting logs.<br />
Diadocidiinae. There is only one genus (Diadocidia) <strong>of</strong> this subfamily in Britain<br />
with three species, two <strong>of</strong> which are found in rotting wood and associated with <strong>the</strong><br />
fungus Peniophora. The larva differs from that <strong>of</strong> o<strong>the</strong>r Myce<strong>to</strong>philidae in having <strong>the</strong><br />
epicranial plates separated ventrally.<br />
Boli<strong>to</strong>philinae. This subfamily has only one genus Boli<strong>to</strong>phila which is Holarctic in<br />
distribution. The known larvae are gregarious and have all been found in <strong>the</strong> s<strong>of</strong>t<br />
fruiting bodies <strong>of</strong> fungi. Boli<strong>to</strong>phila hybrida (Meigen) is very common in Paxillus<br />
involutus (Batsch. ex Fr.) Fr. and B. saundersi Curtis (fig. 81) is usually found in<br />
Hypholomafasciculare (Huds. ex Fr.) Quelet, but both species occur in o<strong>the</strong>r fungi.<br />
Some species appear <strong>to</strong> be restricted <strong>to</strong> a few closely related fungi (i.e. are<br />
stenophagous), e.g. Boli<strong>to</strong>phila cinerea Meigen on Hypholoma and Pholiota spp.<br />
48
Keroplatinae. There are seven <strong>British</strong> genera <strong>of</strong> Keroplatinae.<br />
The larvae <strong>of</strong> this primitive and diverse subfamily spin webs, <strong>the</strong> strands <strong>of</strong> which are<br />
made sticky with drops <strong>of</strong> fluid containing oxalic acid.<br />
Larvae <strong>of</strong> Keroplatus are flattened, somewhat luminous, and live under a large, fiat,<br />
mucilaginous net on <strong>the</strong> underside <strong>of</strong>logs, usually with polyporaceous fungi. They feed<br />
partly on that fungus and partly on o<strong>the</strong>r invertebrates and pupate in a distinct dry<br />
cocoon.<br />
Larvae <strong>of</strong> Orfelia s.s. species are found in turf, grass tussocks, under logs and<br />
bo ulders, in worm tunnels, among mosses and liverworts. Nothing is known <strong>of</strong> <strong>the</strong><br />
biology <strong>of</strong> <strong>the</strong> o<strong>the</strong>r subgenera.<br />
The la rvae <strong>of</strong> Macrocera are poorly known, but have been found in turf, rotting<br />
wood and caves. They are predaceous but some are scavengers at first. The remaining<br />
genera should be sought under logs or rotting wood, though <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
several are unknown.<br />
Sciophilinae. This subfamily contains about one third <strong>of</strong> <strong>the</strong> <strong>British</strong> fauna, and is<br />
divided in<strong>to</strong> five tribes <strong>of</strong> which four are represented in Britain. The <strong>immature</strong> <strong>stages</strong><br />
a re poorly known. The larvae are found on fungi or under bark, inhabit a small<br />
mucilaginous tube supported by threads and, in some species, covered by a dry<br />
irregular, sometimes communal, net.<br />
Sciophila bux<strong>to</strong>ni Freeman occurs in <strong>to</strong>ugh lignicolous polypores, e.g. Coriolus,<br />
Pseudotrametes and Daedaleopsis. Extreme examples <strong>of</strong> polyphagous species are<br />
Sciophi/a lutea Macquart, Rondaniella dimidiata Meigen and Docosia gilvipes Haliday,<br />
larvae <strong>of</strong> <strong>the</strong> last species also being found in <strong>the</strong> nests <strong>of</strong> birds and mammals. Mycomya<br />
species have been bred from fungi associated with dead wood. Larvae <strong>of</strong> Gnoriste are<br />
found in mosses and some Boletina occur in mosses and liverworts as well as <strong>the</strong> more<br />
usual habitats such as rotting wood. Speolepta fep<strong>to</strong>gaster feeds on algae and fungi in<br />
caves (see introduc<strong>to</strong>ry section on caves).<br />
Manotinae. The only <strong>British</strong> species <strong>of</strong> this subfamily is Manota unifurcata<br />
Lundstrom, which is rare and has been reared from a rotting beech with Myxomycetes<br />
(slime mould) but <strong>the</strong> larva is unknown (and <strong>the</strong>refore not included in <strong>the</strong> key).<br />
Myce<strong>to</strong>philinae. Larvae <strong>of</strong> this subfamily are mostly found in s<strong>of</strong>t ground fungi but<br />
some, e.g. Myce<strong>to</strong>phila ornata Stephens, Dyna<strong>to</strong>soma fuscicorne (Meigen), prefer a<br />
wide range <strong>of</strong>lignicolous fungi. O<strong>the</strong>r species are more specialised in <strong>the</strong> range <strong>of</strong> fungi<br />
in which <strong>the</strong>y breed. Myce<strong>to</strong>phi/a a/ea (Laffoon), Exechia nigroscutel/ata Land rock and<br />
Cordyla fasciata Meigen are restricted <strong>to</strong> Russulaceae; Myce<strong>to</strong>phi/a signa<strong>to</strong>ides<br />
Dziedzicki and Exechia separata Lundstrom are restricted <strong>to</strong> Boletaceae. A few species<br />
appear <strong>to</strong> be restricted <strong>to</strong> just one species <strong>of</strong> fungus (i.e. are monophagous) e.g.<br />
Myce<strong>to</strong>phila cingulum Meigen on Polyporus squamosus Fries; Trichonta falcata<br />
Lundstrom on Stereum hirsutum (Willd. ex Fries) and T. apicalis Strobl ( = vernalis<br />
Land rock) on Cafocera cornea (Batsch. ex. Fr.) Fries. Myce<strong>to</strong>phi/a blanda Winnertz is<br />
one <strong>of</strong> <strong>the</strong> few (see Ecology Section) Diptera that induce gall formation in fungi, in this<br />
case on Lactarius deliciosus L. ex Fries.<br />
The slug-like larva (figs 83- 84) <strong>of</strong> Phroniaflavicollis Winnertz ( =strenua Winnertz)<br />
lives on sodden, fallen and barkless branches, where it feeds on moulds. It bears a hard<br />
black conical case (figs 85, 89, 90) made from its own excrement. It leaves this case and<br />
spins a light silky cocoon prior <strong>to</strong> pupation. Phronia braueri Dziedzicki ( = annulata<br />
Winnertz) is found in simila r situations but does not form a case. Epicypta aterrima<br />
(Zetterstedt), on rotten wood, forms a case similar <strong>to</strong> that <strong>of</strong> P.flavicol/is. Some larvae<br />
(<strong>of</strong> this genus) appear <strong>to</strong> live under a mucilaginous 'blob' (fig. 91) without forming a<br />
case. Platurocypta species and Myce<strong>to</strong>phi/a vittipes (Zetterstedt) feed on Myxomycetes<br />
(slime moulds).<br />
49
Sciaridae<br />
(Figs: larvae 87- 88, egg 987, pupa 1126)<br />
The family Sciaridae contains over <strong>10</strong>00 species in over 50 genera and is represented<br />
in most regions <strong>of</strong> <strong>the</strong> world. In Britain about <strong>10</strong>0 species in 18 genera have so far been<br />
recorded but o<strong>the</strong>rs are now being discovered following a recent Handbook <strong>to</strong> this longneglected<br />
family (Freeman, 1983, 1987). For a long time sciarids were regarded as<br />
comprising a subfamily <strong>of</strong> <strong>the</strong> Myce<strong>to</strong>philidae but <strong>the</strong>y are now given family status.<br />
The larvae are white with a sharply contrasting shiny black head and remarkably<br />
uniform general appearance (figs 87- 88). The terminal abdominal segment is slightly<br />
lobate and is used as a leg in locomotion. There are four larval instars, <strong>the</strong> first<br />
metapneustic, <strong>the</strong> second and third propneustic and <strong>the</strong> fourth hemipneustic (see fig. I).<br />
Diagnostic features are given in <strong>the</strong> key <strong>to</strong> families.<br />
The larvae are usually associated with decaying vegetable matter, excrement, rotten<br />
wood and fungi .<br />
Three species are recorded as pests <strong>of</strong> mushrooms: Lycoriella so/ani (Winnertz), L.<br />
auripi/a (Winnertz) and Bradysia brunnipes (Meigen). The larvae tunnel in<strong>to</strong> <strong>the</strong><br />
mushroom <strong>to</strong> feed, but more damage appears <strong>to</strong> be caused through <strong>the</strong> transmission <strong>of</strong><br />
disease and mites associated with <strong>the</strong> bacterial breakdown <strong>of</strong> <strong>the</strong> fungi (Hussey, Read<br />
& Hesling, 1969). Larvae may still be recognizable after reconstitution in dehydrated<br />
mushroom soup!<br />
O<strong>the</strong>r species <strong>of</strong> economic importance through larvae damaging plants and seedlings<br />
in greenhouses include Bradysia amoena Winnertz, B. brunnipes, B. paupera<br />
Tuomikoski, B. tritici (Coquillett) (<strong>the</strong> mossfty <strong>of</strong> orchid growers), Ctenosciara<br />
hyalipennis (Meigen), Lycoriella auripila and Pnyxia scabei (Hopkins) (on cucumbers,<br />
Speyer, 1922).<br />
Larvae <strong>of</strong> Plas<strong>to</strong>sciara and Xylosciara are usually found under bark or in rotting<br />
wood, but P. perniciosa (Edwards) is a common greenhouse pest, destroying <strong>the</strong> roots<br />
and stems <strong>of</strong> cucumber seedlings. Adults <strong>of</strong> this species have also occurred in numbers<br />
indoors, possibly due <strong>to</strong> larvae feeding in decaying peat, used in <strong>the</strong> insulating lining <strong>of</strong><br />
old fashioned steel safes. Adults indoors may also originate from larvae feeding in<br />
overdamp compost around potted plants.<br />
The larvae <strong>of</strong> several species have been recorded from old nests <strong>of</strong> birds and<br />
mammals (Hicks, 1959- 1971; Hackman, 1963) e.g. Bradysia spp., Corynoptera spp.,<br />
Lycoriella sagraria (Felt), L. lundstromi (Frey), Trichosia basdeni Freeman, T.<br />
coarctata Winnertz. Most nests contain droppings and o<strong>the</strong>r decaying material which<br />
may attract <strong>the</strong>m and <strong>of</strong> course some are lined with a mixture <strong>of</strong> mud and dung.<br />
Occasionally mass movements <strong>of</strong> sciarid larvae have been reported in Europe, <strong>the</strong><br />
larvae moving in columns 8- 25 cm wide and 2- 3 metres long. The species involved are<br />
Sciara thomae (L.) and S. militaris Nowicki. The former occurs fairly commonly in<br />
Britain, but <strong>the</strong> phenomenon has not yet been observed here and its cause or purpose is<br />
quite unknown.<br />
There is plenty <strong>of</strong> scope for research on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> this family. Thomas<br />
( 1930) describes <strong>the</strong> larva and pupa <strong>of</strong> Bradysia aprica (Winnertz) (as Sciara nitidicollis,<br />
see Freeman, 1983) from decaying celery roots. Mad war (1934) describes <strong>the</strong> larvae <strong>of</strong><br />
Pnyxia scabiei and (1937) <strong>of</strong> Lycoriella auripila, Heterosciara semialata (Edwards),<br />
Bradysiafenestralis (Zetterstedt) and Plas<strong>to</strong>sciara perniciosa. Very little has been done<br />
since.<br />
Sca<strong>to</strong>psidae<br />
(Figs: larvae 92- <strong>10</strong>0, pupae, 1127- 1128)<br />
The Sca<strong>to</strong>psidae is a little known, little worked family. About 300 species have been<br />
described, contained in 18 genera. Four subfamilies are recognised but larvae <strong>of</strong> only<br />
50
two <strong>of</strong> <strong>the</strong>se are known, so a key cannot be given and when more larvae are known even<br />
<strong>the</strong> family key may need emendation. All four subfamilies occur in Britain and such<br />
biological information as is available is given under <strong>the</strong>se.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> very few species are known. Of <strong>the</strong> 37 <strong>British</strong> species (in 15<br />
genera) <strong>the</strong> larvae <strong>of</strong> only four have been adequately described.<br />
Aspistinae. The <strong>immature</strong> <strong>stages</strong> are unknown. Adults <strong>of</strong> <strong>the</strong> only <strong>British</strong> species,<br />
Aspistes berolinensis Meigen, are rare and found in sandy places. In Britain <strong>the</strong> species<br />
has so far only been found in coastal sand dunes but continental records are from far<br />
inland. In Czechoslovakia adults have been reared from 'sifted litter' where it is<br />
thought <strong>the</strong>re may be two generations a year.<br />
Ectaetinae. Laurence ( 1953) describes <strong>the</strong> larva <strong>of</strong> Ectaetia platyscelis Loew (figs 96,<br />
I 00) which he reared from a deposit <strong>of</strong> wet orange coloured detritus lying in a hole in a<br />
lime tree trunk. Adults <strong>of</strong> this and <strong>the</strong> o<strong>the</strong>r two <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus (whose<br />
larvae remain unknown) have been reared from rotting trees.<br />
Psectrosciarinae. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Psectrosciarinae are unknown.<br />
The subfamily is represented in Britain by three species <strong>of</strong> <strong>the</strong> genus <strong>An</strong>apausis.<br />
<strong>An</strong>apausis so/uta (Loew) is common and widespread in <strong>the</strong> adult stage and is frequently<br />
found swarming in large number on fences and poles; it may also occur indoors where I<br />
have seen specimens that were found in cracks behind a kitchen sink. A. talpae Verrall<br />
was discovered swarming on a willow trunk.<br />
Sca<strong>to</strong>psinae. The larva <strong>of</strong> Sca<strong>to</strong>pse notata (L.) and Coboldiafuscipes (Meigen) have<br />
been found in a variety <strong>of</strong> decaying plant and animal materials, e.g. wasps' nests, bulbs<br />
and onions, green ginger damaged by water (Lyall, 1929), wastes from fruit and wine<br />
canneries, excrement and decomposing fungi. The larva <strong>of</strong> S. notata (figs 92, 97),<br />
described by Morris (1918), is somewhat flattened dorso-ventrally and dorsally has<br />
thick setae, arranged in a regular pattern, which are usually full <strong>of</strong> detritus when <strong>the</strong><br />
larva is first found. It is peripneustic with one pair <strong>of</strong> prothoracic and eight pairs <strong>of</strong><br />
abdominal spiracles. The larva <strong>of</strong> Coboldia is similar but <strong>the</strong> dorsal hairs are not<br />
arranged in a regular pattern (fig. 94). Bovien (1935) found that in C.fuscipes <strong>the</strong> length<br />
<strong>of</strong> <strong>the</strong> caudal spiracular processes varied with <strong>the</strong> degree <strong>of</strong> dampness <strong>of</strong> <strong>the</strong> breeding<br />
medium. In dry conditions <strong>the</strong>y were short, but in wet conditions <strong>the</strong>y were long and<br />
enabled <strong>the</strong> larvae <strong>to</strong> hang below <strong>the</strong> meniscus (like some o<strong>the</strong>r aquatic Nema<strong>to</strong>cera).<br />
The larva <strong>of</strong> Rhexosa has not yet been found in Britain but American species (fig. 93)<br />
have been found in <strong>the</strong> decaying bark <strong>of</strong> dead or dying deciduous trees. Lyall ( 1929)<br />
gives notes on R. subnitens (Verrall).<br />
Apilosca<strong>to</strong>pse species have been reared from fungi: A . jfavicollis (Meigen) from<br />
Tricholoma pessundatum Quelet and A. scutellata (Loew) from Bjerkandera adusta.<br />
Parasca<strong>to</strong>pse li<strong>to</strong>rea (Edwards) (figs 95, 98) is a coastal species in Britain but <strong>the</strong><br />
interesting larva has only recently been discovered and described from saltings in<br />
Poland (Szadziewski, 1979). The larvae were found in numbers (April-July) from salt<br />
soil covered with Salicornia patula Duvai-Jouve and Aster tripolium L. Probably <strong>the</strong><br />
larvae <strong>of</strong> both <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus are <strong>to</strong> be sought in salt-marshes.<br />
Adults <strong>of</strong> Holoplagia transversalis (Loew) have occasionally been found with ants<br />
(Lasius fuliginosus Latreille) and H. richardsi Edwards has been reared from rotting<br />
elm, but <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are unknown.<br />
Adults <strong>of</strong> Reichertella geniculata (Zetterstedt) have been found congregated on a<br />
poppy seed-head but <strong>the</strong> life-his<strong>to</strong>ry is unknown. Wehrmeister (1924) figures <strong>the</strong> anal<br />
papillae (fig. 99) <strong>of</strong> R. pulicaria Loew but gives no habitat details.<br />
Clearly, careful rearing from <strong>the</strong> habitats mentioned, especially rotting trees, should<br />
elucidate new life-his<strong>to</strong>ries in this neglected family.<br />
51
Cecidomyiidae<br />
(Figs: larvae <strong>10</strong>1- 122, 125; eggs 989- 990; pupae 123-124, 1129-1130;<br />
mines and galls 1289-1290, 1311- 1315)<br />
About 4500 species <strong>of</strong> Cecidomyiidae (gall midges) have so far been described<br />
throughout <strong>the</strong> world. Three subfamilies are recognised, all <strong>of</strong> which occur in Britain,<br />
and <strong>the</strong> larvae may be distinguished as follows.<br />
<strong>An</strong>us slit-like (fig. <strong>10</strong>2) and usually ventral (dorsal in some predaceous species, e.g.<br />
Les<strong>to</strong>dip/osis) . . . . . . . . . . . . . . . . . . . . . . 2<br />
<strong>An</strong>us rqunded, terminal (fig. <strong>10</strong>1) . . . . . . . . . . . . . . . . 3<br />
2 Abdominal segment 8 with four dorsal papillae (fig. <strong>10</strong>3, dp), abdominal segments 1- 7 with<br />
four posteroventral papillae (pv) (sometimes imperceptible); not gall-forming . .<br />
. . . . . . . . . . . . . . . . . . Porricondylinae (pa rt)<br />
Abdominal segment 8 with two dorsal papillae; abdominal segments 1- 7 with 2<br />
posteroventral papillae; some are gall-forming . . . . . . . . Cecidomyiinae<br />
3 Lateral papillae in groups <strong>of</strong> 2; anus usually protrusible; not gall-forming Lestremiinae<br />
Lateral papillae in groups <strong>of</strong> 3 (fig. <strong>10</strong>3, lp) anus non-protrusible; not gall-forming . .<br />
. . . . . . . . . . . . . . . . . Porricondylinae (Heteropezini)<br />
Cecidomyiid larvae are <strong>of</strong>ten brightly coloured red, orange, pink, yellow or sometimes<br />
white. The distinctive sterna! spatula or 'breastbone' (figs <strong>10</strong>3-116) is present<br />
only in final (third) ins tar larvae and is absent in some species (e.g. Contarinia geniculati<br />
Reuter, a salmon pink larva in <strong>the</strong> florets <strong>of</strong> meadow fox tail and cocksfoot grasses) and<br />
in paedo!;enetic larvae (e.g. Mias<strong>to</strong>r, Mycophila).<br />
The spatula is best developed in soil-dwelling larvae where it is <strong>of</strong> use in tunnelling<br />
and excavating a pupal chamber. This can cause wear <strong>of</strong> <strong>the</strong> spatula and caution is <strong>the</strong>n<br />
necessary in identification from this feature alone (Milne, 1961 ).<br />
Eggs are usually small and ovoid and about <strong>10</strong>0 are produced by <strong>the</strong> adult females,<br />
except in paedogenetic forms, when <strong>the</strong>y are few and large. Paedogenesis is more fully<br />
discussed in <strong>the</strong> introduc<strong>to</strong>ry section on Immature Stages (p. 24).<br />
There is still much <strong>to</strong> be done on <strong>the</strong> larvae <strong>of</strong> Cecidomyiidae, particularly on <strong>the</strong>ir<br />
description and detailed systematics. The series <strong>of</strong> books by H. F. Barnes (1946-1956)<br />
and Nijveldt ( 1969) is a mine <strong>of</strong> information and more specialised works are cited<br />
under each subfamily. Useful studies <strong>of</strong> gall midge communities, <strong>the</strong>ir life-his<strong>to</strong>ries<br />
and parasites are provided by Askew & Ruse (1974, on birch leaves), Hodges (1969,<br />
birch catkins), Milne (1960, clover), Otter (1938, knapweed), Parnell (1963, broom),<br />
and I. J. Wyatt (1964, mushrooms).<br />
Lestremiinae. Larvae <strong>of</strong> this subfamily are poorly known but have been found in<br />
decaying wood and vegetation, plant wounds and fungi, including mushrooms.<br />
Larvae <strong>of</strong> <strong>the</strong> mushroom infesting species are described and keyed by Wyatt ( 1964)<br />
(e.g. Lestremia cinerea, Mycophila speyeri, M . barnesi). Mycophila species are usually<br />
paedogenetic (i.e. with reproducing larvae, see Introduction and Porricondylinae<br />
below). The sterna! spatulae (<strong>of</strong> <strong>the</strong> sexual cycle) <strong>of</strong> two <strong>of</strong> <strong>the</strong> mushroom pest species<br />
are illustrated (figs 113- 114). Monardia ulmaria Edwards has been reared from rotten<br />
elm. The reddish larvae <strong>of</strong> Campylomyza ormerodi (Kieffer) (fig. 1 05) have been alleged<br />
<strong>to</strong> attack <strong>the</strong> tap root <strong>of</strong> red clover at ground level and <strong>the</strong> plant apex but this observation<br />
requires confirmation; C. pumila Winnertz has been reared from an Ascomycete<br />
from <strong>the</strong> United States. Adults <strong>of</strong> Aprionus species occur around old logs and stumps,<br />
especially beech. Peromyia species occur under bark and leaf litter and P. fungicola<br />
(Kieffer) (fig. 120) has been reared from a mould growing on <strong>the</strong> surface <strong>of</strong> a <strong>to</strong>ads<strong>to</strong>ol<br />
(Lactarius).<br />
Porricondylinae. Larvae <strong>of</strong> this subfamily are myce<strong>to</strong>phagous in decaying vegetation<br />
and wood and may be secondary invaders in <strong>the</strong> living tissues <strong>of</strong> higher plants.<br />
52
In his keys <strong>to</strong> mushroom infesting cecid larvae Wyatt (1964) includes Heteropeza<br />
pygmaea Winnertz and Henria psa/liotae Wyatt, which are paedogenetic. He has also<br />
reviewed <strong>the</strong> phenomenon <strong>of</strong> paedogenesis in <strong>the</strong> family (Wyatt 1967) including <strong>the</strong><br />
well known larvae <strong>of</strong> Mias<strong>to</strong>r which are fairly easily found in numbers under <strong>the</strong> bark<br />
<strong>of</strong> rotting birch where <strong>the</strong>y <strong>of</strong>ten form star-shaped clusters. The mo<strong>the</strong>r larvae increase<br />
in size merely by distention <strong>of</strong> <strong>the</strong> cuticle and <strong>the</strong> daughter larvae can be clearly seen<br />
inside (fig. 118). The striking larvae <strong>of</strong> Parepidosis (fig. 117) are found under bark and<br />
leaf litter.<br />
Panel ius ( 1965) covers <strong>the</strong> subfamily with notes on larvae and biology where known.<br />
Mohn ( 1955) gives detailed descriptions <strong>of</strong> larvae.<br />
Cecidomyiinae. This subfamily contains <strong>the</strong> many gall-producing forms which earn<br />
<strong>the</strong> family its common name. Not all form galls however; some feed in <strong>the</strong> stems<br />
and flower heads <strong>of</strong> plants, o<strong>the</strong>rs are myce<strong>to</strong>phagous and some are preda<strong>to</strong>rs or<br />
parasi<strong>to</strong>ids. Larvae <strong>of</strong> <strong>the</strong> tribe Oligotrophidi mostly live freely in flower heads or cause<br />
leaf rolls or swollen stems, but Craneiobia corni (Giraud) causes prominent galls on<br />
dogwood (fig. 1313).<br />
The large genus Dasineura ( 136 <strong>British</strong> species) contains many species <strong>of</strong> economic<br />
importance (see table in Economic Section). The white larvae <strong>of</strong> D. brassicae (fig. <strong>10</strong>9)<br />
cause <strong>the</strong> 'bladder pod' condition which prevents normal seed development in cabbage,<br />
swede, turnip, rape and radish and has increased in importance with <strong>the</strong> increased<br />
growth <strong>of</strong> oil seed rape. D. leguminicola (fig. 11 0) attacks <strong>the</strong> inflorescences <strong>of</strong> red and<br />
white clover; <strong>the</strong> larvae are yellow at first, <strong>the</strong>n become salmon pink. D. trifolii (fig. Ill)<br />
forms galls on <strong>the</strong> leaves <strong>of</strong> clover plants which cause <strong>the</strong> leaflets <strong>to</strong> stick <strong>to</strong>ge<strong>the</strong>r; <strong>the</strong><br />
larvae are white at first and later become orange. The red or orange larvae <strong>of</strong> D.<br />
alopecuri (fig. 112) feed and pupate in <strong>the</strong> florets <strong>of</strong> cocksfoot grass (Dactylis glomerata<br />
(L.)). The galls <strong>of</strong> D. ulmaria (Bremi) may be common on meadow sweet (Spiraea).<br />
Mayetiola destruc<strong>to</strong>r (Say), <strong>the</strong> well known hessian fly (figs 116, 122, 1311), attacks<br />
<strong>the</strong> straw <strong>of</strong> wheat and causes widespread damage throughout <strong>the</strong> wheat growing areas<br />
<strong>of</strong> N. Africa, <strong>the</strong> U.S.A., New Zealand and part <strong>of</strong> Europe, but fortunately is less<br />
serious in Britain. The pupae (figs 123, 1130) are formed inside <strong>the</strong> last larval skin<br />
(which is more characteristic <strong>of</strong> Cyclorrhapha - see introduc<strong>to</strong>ry section on pupae)<br />
and are sometimes called 'flax seeds' because <strong>of</strong> <strong>the</strong>ir superficial resemblance <strong>to</strong> such.<br />
Rhabdophaga species (26 in Britain) (fig. 1315) are mostly associated with willows.<br />
Rondaniola bursaria (Bremi) forms <strong>the</strong> galls frequently seen on ground ivy ( Glechoma<br />
hederacea L.) (fig. 1314).<br />
Sylven ( 1975) deals with larval structures and habits <strong>of</strong> Oligotrophidi.<br />
True galls, mostly on flowers and buds but also on leaves, are induced by larvae <strong>of</strong> <strong>the</strong><br />
super-tribe Asphondyiidi.<br />
The larvae <strong>of</strong> Lasiopterini and Oligotrophini mainly cause simple galls on twigs,<br />
petioles, midribs or leaves, a few occur in <strong>the</strong> seeds <strong>of</strong> Compositae and at least one<br />
species <strong>of</strong> Brachyneurini, Brachyneurina peniophorae Harris, forms galls in <strong>the</strong> fungus<br />
Peniophora cinerea (Fr.) Cook (Harris & Evans, 1979).<br />
Members <strong>of</strong> <strong>the</strong> largest supertribe, <strong>the</strong> Cecidomyiidi, have a very wide range <strong>of</strong><br />
habits, including free living, phy<strong>to</strong>phagous and mycophagous forms, gall makers,<br />
preda<strong>to</strong>rs and internal parasi<strong>to</strong>ids. The large genus Contarinia (72 <strong>British</strong> species)<br />
contains most <strong>of</strong> <strong>the</strong> species <strong>of</strong> economic importance (see economic section and Harris,<br />
1966), <strong>the</strong> larvae <strong>of</strong> which develop in malformed flowers <strong>of</strong> various Dicotyledons,<br />
especially Leguminosae (e.g. C. pisi on peas) (fig. <strong>10</strong>7); o<strong>the</strong>rs develop in <strong>the</strong> spikelets <strong>of</strong><br />
Gramineae (e.g. C. tritici on wheat (figs <strong>10</strong>8, 1312) (see also Si<strong>to</strong>diplosis below), and<br />
C. merceri Barnes and C. geniculati (Reuter) on Alopecurus). O<strong>the</strong>rs are found in galls<br />
on trees or form galls in <strong>the</strong> leaves (e.g. Pu<strong>to</strong>niella pruni Kaltenbach on plum) and<br />
inflorescences <strong>of</strong> Monocotyledons and Dicotyledons, and a few attack developing fruit<br />
(e.g. Contarinia pyrivora Riley on pear).<br />
53
Cecidomyia species are associated with conifers. C. pini (fig. 121) larvae feed in resin<br />
masses on a number <strong>of</strong> species <strong>of</strong> Coniferae.<br />
Myocecis ova/is Edwards induces a localized hypha! mat in <strong>the</strong> fungus Hypoxylon<br />
rubiginosum Pers ex Fr. (Evans, 1970). The bright red and orange larvae <strong>of</strong><br />
Mycodiplosis species feed on rusts and mildews on a wide range <strong>of</strong> plants, including<br />
many agricultural crops. The commonest <strong>British</strong> species, M . saundersi Barnes, feeds on<br />
a rust on creeping thistle (Cirsium arvense).<br />
Les<strong>to</strong>diplosis (fig. 119) is ano<strong>the</strong>r large genus (34 <strong>British</strong> species) found in a wide<br />
range <strong>of</strong> microhabitats. The extremely active larvae are predaceous on mites, certain<br />
beetle larvae and on o<strong>the</strong>r cecidomyiids. Aphidoletes and Monobremia larvae are<br />
specialized preda<strong>to</strong>rs on aphids and A . aphidimyza (Rondani) (fig. 125) has been used<br />
as an effective biocontrol agent (Harris, 1982).<br />
The larva <strong>of</strong> Endopsylla agilis de Meijere is an endoparasite <strong>of</strong>Psyllidae (Horn.) and<br />
that <strong>of</strong> Endaphis perjidus Kieffer is an endoparasite <strong>of</strong> aphids.<br />
Si<strong>to</strong>diplosis mosellana (Gehin) is ano<strong>the</strong>r wheat blossom midge <strong>the</strong> larva <strong>of</strong> which<br />
can be distinguished by its colour (orange <strong>to</strong> red) from that <strong>of</strong> Contarinia tritici<br />
(yellow- see above) and <strong>the</strong> relative positions occupied in <strong>the</strong> wheat flower (fig. 1312).<br />
Mohn ( 1955) gives detailed descriptions <strong>of</strong> larvae <strong>of</strong> Cecidomyiinae.<br />
54
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Brachycera<br />
Posterior spiracles close <strong>to</strong>ge<strong>the</strong>r and more or less concealed within a terminal fissure on <strong>the</strong><br />
anal segment . 2<br />
Posterior spiracles widely separated on <strong>the</strong> anal, penultimate or antepenultimate abdominal<br />
segment and not concealed (even if in a fissure) . 4<br />
2 Terminal fissure vertical; head capsule not strongly sclerotized, and retractile; body cylindrical<br />
with smooth cuticle; anterior seven segments all, or mostly, encircled by a row <strong>of</strong><br />
projections which sometimes bear apical spicules a nd function as pro legs. Mainly in moist<br />
or wet soils, Sphagnum, etc.; inactive (figs 157- 160) . Tabanidae (p. 60)<br />
Terminal fissure transverse; head capsule strongly sclerotized, and non-retractile; body<br />
depressed with roughened cuticle 3<br />
3 <strong>An</strong>al slit on <strong>the</strong> central part <strong>of</strong> <strong>the</strong> anal segment bordered with teeth-like projections and a<br />
row <strong>of</strong> similar projections arranged transversely in front <strong>of</strong> <strong>the</strong> anal slit (fig. 141 ); thoracic<br />
tergites with a more or less well defined smooth area on <strong>the</strong> median part <strong>of</strong> each tergite;<br />
amphipneustic; under bark <strong>of</strong> decaying trees; inactive (figs 141- 143) Xylomyidae (p. 58)<br />
<strong>An</strong>al slit area without such teeth-like projections; thoracic tergites uniformly roughened<br />
(shagreened like shark-skin); holo- or peripneustic; inactive; terrestrial species broad a nd<br />
flattened (figs 126--130), in woodland debris, dung, compost heaps and o<strong>the</strong>r decaying<br />
vegetable material; aquatic species <strong>of</strong>ten having a tapering abdomen terminating in a<br />
respira<strong>to</strong>ry siphon (figs 131-140). Stratiomyidae (p. 57)<br />
4 Posterior spiracles on <strong>the</strong> last apparent body segment . 5<br />
Posterior spiracles anterior <strong>to</strong> <strong>the</strong> last body segment . 11<br />
5 Head capsule very short and minute; body s<strong>of</strong>t, broad, pear-shaped with enlarged abdomen;<br />
last abdominal segment without processes but with two distinct spiracles; anterior spiracle<br />
very small; inactive, endoparasites <strong>of</strong> spiders (<strong>the</strong> early stage larvae are free-living, active<br />
triungulins, elongated with long setae) (figs 177- 182) . Acroceridae (p. 64)<br />
Head capsule prominent; last abdominal segment usually with processes; free living . 6<br />
6 Head and one or more thoracic tergites strongly sclerotized; posterior spiracles on a<br />
sclerotized plate with projecting processes; under bark <strong>of</strong> decayed trees, sometimes within<br />
old burrows; inactive (fig. 144) . . Xylophagidae (p. 58)<br />
Head not strongly sclerotized; posterior spiracles not on a sclerotized plate . 7<br />
7 <strong>An</strong>al segment with two or four, usually pointed, fleshy lobes open at end in a concave cleft<br />
containing <strong>the</strong> posterior spiracles 8<br />
<strong>An</strong>al segment o<strong>the</strong>rwise 9<br />
8 Lobes <strong>of</strong> anal segment subequal in size; soil dwelling (probably at least partially predaceous)<br />
(figs 145- 151) . Rhagionidae(p. 59)<br />
Lobes <strong>of</strong> anal segment with ventral pair longer and more strongly developed (fig. 206);<br />
mouthparts with metacephalic rods (upper rods) swollen or spatulate at tip (fig. 215);<br />
predaceous in damp soil, under bark, in decaying wood, some in wet sand on sea shore<br />
(figs 206--238) . Dolichopodidae (p. 67)<br />
9 <strong>An</strong>al segment rounded at end with lateral furrows evident and sometimes a small median<br />
projection below <strong>the</strong> posterior spiracles (fig. 196); (or, Ocydromia, lacking lateral furrows<br />
and with a distinct lobe below spiracles in dung, fig. 193); terrestrial, predaceous (figs<br />
189- 205) Empididae (part) (p. 65)<br />
<strong>An</strong>al segment usually with long float hairs, borne on obvious fleshy lobes; aquatic,<br />
predaceous . I 0<br />
<strong>10</strong> Larvae slightly flattened dorsoventrally; slender lateral and dorsolateral tubercles <strong>of</strong> progressively<br />
increasing size on abdominal segments 1- 7; prolegs on all 8 abdominal<br />
segments; two large caudal tubercles fringed with hairs on terminal segment (figs 152- 156)<br />
A<strong>the</strong>ricidae (p. 59)<br />
Larvae not dorsoventrally flattened; no lateral or dorsolateral tubercles on segments 1- 7;<br />
pro legs on 7 or 8 <strong>of</strong> <strong>the</strong> abdominal segments; tubercles on terminal segment smaller (figs<br />
201- 205) Empididae (part) (p. 65)<br />
11 Posterior spiracles on <strong>the</strong> apparent penultimate segment; abdominal segments not subdivided,<br />
<strong>the</strong> body consisting <strong>of</strong> 11 or 12 apparent segments, exclusive <strong>of</strong> <strong>the</strong> head . 12<br />
Posterior spiracles on <strong>the</strong> apparent antepenultimate segment; abdominal segments 1-6<br />
sub-divided, <strong>the</strong> body thus apparently consisting <strong>of</strong> up <strong>to</strong> 20 segments, exclusive <strong>of</strong> <strong>the</strong><br />
head 13<br />
55
12 Thoracic segments each with two long setae, one on each side <strong>of</strong> ventrolateral margin; anal<br />
segment with six or more long setae; free living in soil or decayed wood; predaceous,<br />
ec<strong>to</strong>parasitic (figs 165-176) Asilidae (p. 61)<br />
Thoracic segments without, or with only weak setae; anal segment without setae; preda<strong>to</strong>rs,<br />
parasites or inquilines in nests <strong>of</strong> bees and wasps or in egg cases <strong>of</strong> grasshoppers; resembling<br />
a bee or wasp grub; inactive (young larvae are slender, active, free-living triungulins<br />
with a long slender seta on each side <strong>of</strong> each thoracic segment and two very long setae on<br />
<strong>the</strong> anal segment, fig. 183) (figs 183- 188) . Bombyliidae (p. 64)<br />
13 Dorsal extension <strong>of</strong> head capsule extending in<strong>to</strong> <strong>the</strong> thorax from <strong>the</strong> head posteriorly, with a<br />
spatulate apex (fig. 162); head capsule with two ventral projections; in soil (<strong>of</strong>ten sandy);<br />
predaceous, active with snake-like movement or 'jumps' (figs 161- 162) .<br />
Therevidae (p. 63)<br />
Dorsal extension <strong>of</strong> head capsule extending in<strong>to</strong> <strong>the</strong> thorax from <strong>the</strong> head posteriorly not<br />
spatulate at apex (fig. 163); head capsule without ventral projections; predaceous on<br />
insects in household s<strong>to</strong>red foods, wood or fungi, active with movements as Therevidae<br />
(figs 163- 164) . . Scenopinidae (p. 63)<br />
56
Notes on families <strong>of</strong> Brachycera<br />
Stratiomyidae<br />
(Figs: larvae 126-140, eggs 991-992, emergence <strong>of</strong> adult 1330)<br />
The family Stratiomyidae (soldier flies) contains some 2300 species <strong>of</strong> which 50<br />
species in 12 genera occur in Britain. Six subfamilies are recognised, all <strong>of</strong> which are<br />
represented in Britain and distinguishable in <strong>the</strong> larval stage as follows (based on<br />
Rozkosny, 1982).<br />
Last abdominal segment usually rectangular or elongated and tubelike, with a coronet <strong>of</strong><br />
pinnate or plumose float-hairs at <strong>the</strong> apex. If <strong>the</strong> apical coronet is absent, <strong>the</strong>n ei<strong>the</strong>r <strong>the</strong><br />
dorsal abdominal setae are in two transverse rows or <strong>the</strong> abdominal setae are flat with a<br />
frayed margin . . . . . . . . . . . . . . . . . . . . . . . 2<br />
Last abdominal segment rounded posteriorly, without a coronet <strong>of</strong> float hairs at <strong>the</strong> apex.<br />
Dorsal and ventral abdominal setae arranged in one transverse row; never flat with frayed<br />
margin . . . . . . . . . . . . . . . . . . . . . . . . 3<br />
2 Larvae with an apical coronet <strong>of</strong> setae (figs 132- 133, 135, 139- 140) . . Stratiomyinae<br />
Larvae without an apical coronet <strong>of</strong> setae (fig. 13 I); terrestrial, among semi-aquatic mosses<br />
on s<strong>to</strong>nes etc . . . . . . . . . . . . . . . . . . . Clitellariinae<br />
3 Body setae surrounded by bristle-like hairs which form more or less distinct tufts; anal<br />
segment fringed with fine setae (fig. 126). Terrestrial . . . . . . . . Beridinae<br />
Body setae not surrounded by bristle-like hairs; anal segment without marginal fringe 4<br />
4 Large larvae (18- 20 mm long) with a dense covering <strong>of</strong> adpressed yellow hairs; posterior<br />
spiracular opening almost apical (fig. I 30) . . . . . . . . . . . Hermetiinae<br />
Smaller larvae (at most 12 mm long), with surface-hairs sparse or absent; posterior<br />
spiracular opening dorsal . . . . . . . . . . . . . . . . 5<br />
5 Medium sized larvae, with distinct ocular lobes (ol) (figs 128- 129). Terrestrial . Sarginae<br />
Small larvae (4·5-7·5 mm long), without ocular lobes (fig. 127). Under bark . . . .<br />
. . . . . . . . . . . . . . . . . . . . . . . . Pachygastrinae<br />
Keys <strong>to</strong> genera and species <strong>of</strong> larvae and puparia <strong>of</strong> European Stratiomyidae are<br />
provided by Rozkosny (1982, 1983), who also gives much biological information.<br />
Brindle ( 1959) provides a useful key <strong>to</strong> genera, including those now placed in o<strong>the</strong>r<br />
families. The pupae <strong>of</strong> Stratiomyidae are formed inside <strong>the</strong> last larval skin (see<br />
introduc<strong>to</strong>ry section on pupae).<br />
Beridinae. The larvae <strong>of</strong> Beris (fig. 126) live in compost, decaying vegetation, wet<br />
moss, etc. and B.fuscipes Meigen is <strong>the</strong> only <strong>British</strong> species unknown in this stage. The<br />
only o<strong>the</strong>r <strong>British</strong> genus is Chorisops with two species. The larvae <strong>of</strong> C. tibialis (Meigen)<br />
has been found among dead leaves and a puparium <strong>of</strong> C. naga<strong>to</strong>mii Rozkosny (1982,<br />
described since <strong>the</strong> Diptera Check List) was found in refuse washed up on a muddy<br />
stream bank (water meadows by River Test, P.J. Chandler).<br />
Clitellariinae. Larvae <strong>of</strong> <strong>the</strong> reputedly <strong>British</strong> Clitellaria ephippium (F.) have been<br />
recorded on several occasions by continental authors in nests <strong>of</strong> <strong>the</strong> ant Lasius<br />
fuliginosus (Latreille) and also at <strong>the</strong> roots <strong>of</strong> a walnut tree and in forest soil. The larval<br />
stage appears <strong>to</strong> last 3 or 4 years.<br />
Three <strong>of</strong> <strong>the</strong> four <strong>British</strong> species <strong>of</strong> Nemotelus (fig. 131) have been described in <strong>the</strong><br />
larval stage by Brindle (1964b). The larvae occur in or on <strong>the</strong> mud <strong>of</strong> marshes and at<br />
least N. uliginosus (L.) and N. notatus Zetterstedt are found in estuarine marshes or<br />
inland in brine pits and similar habitats.<br />
Oxycera larvae (fig. 132) occur in hygropetricous habitats, i.e. in mosses growing in<br />
water flowing down inclined rocks or in semi-aquatic mosses in marshes. All but two<br />
(0. dives Loew and 0 . terminata Meigen) are keyed in <strong>the</strong> larval stage by Brindle<br />
(1964b). (0. nigripes Verrall is synonymized with 0. pygmaea (Fallen) by Rozkosny,<br />
1983).<br />
57
The larva <strong>of</strong> Vanoyia tenuicornis (Meigen) is unknown but should be sought along<br />
streams and marshes (including salt marshes) where <strong>the</strong> adults occur (<strong>of</strong>ten on <strong>the</strong><br />
leaves <strong>of</strong> Ligustrum).<br />
Pachygastrinae. Larvae <strong>of</strong> this subfamily (fig. 127) are found under <strong>the</strong> bark <strong>of</strong><br />
decaying trees. Some species appear <strong>to</strong> be associated with particular species <strong>of</strong> trees<br />
and <strong>the</strong>y would be a worthy subject for detailed investigation, e.g. Neopachygaster<br />
meromelaena (Dufour) ( = P. orbitalis Wahlberg) (holly, willow); Pachygaster atra<br />
(Panzer) (elm); Zabrachia ( = Pachygaster) minutissima Zetterstedt (pine); Eupachygaster<br />
tarsalis Zetterstedt (beech, poplar) and Praomyia leachii Curtis (oak). Brindle<br />
(1962b, 1966) keys all <strong>the</strong> <strong>British</strong> species.<br />
Sarginae. The larva <strong>of</strong> <strong>the</strong> common Chloromyiaformosa (Scopoli) is found in dung.<br />
Microchrysa polita (L.) (fig. 128) occurs in dung, grass-heaps, compost, garden refuse,<br />
under moss on old tree trunks, etc., and is <strong>the</strong> most commonly encountered larva <strong>of</strong> <strong>the</strong><br />
family. M.flavicornis (Meigen) has been found in soil under moss on a tree trunk and<br />
may share <strong>the</strong> wide range <strong>of</strong> habitats favoured by M . polita but M.jiavicornis is found<br />
in dung (pig, cow, horse).<br />
Four <strong>of</strong> <strong>the</strong> six <strong>British</strong> species <strong>of</strong> Sargus are known in <strong>the</strong> larval stage from dung and<br />
decaying grass heaps. Roberts ( 1969) has studied <strong>the</strong> structure <strong>of</strong> <strong>the</strong> larval mouth parts<br />
<strong>of</strong> Sargus and concludes that it is a detritus feeder.<br />
Brindle (1965e) keys <strong>the</strong> larvae <strong>of</strong> this subfamily.<br />
Stratiomyinae. The larvae <strong>of</strong> this subfamily are all aquatic and live in shallow<br />
standing water and in ponds or marshes, where <strong>the</strong>y feed on microscopic organisms.<br />
Brindle (1964a) keys three <strong>of</strong> <strong>the</strong> <strong>British</strong> species <strong>of</strong> Stratiomys (figs 133- 138) and<br />
Rozkosny (1982) includes all four.<br />
A key <strong>to</strong> larvae <strong>of</strong> four <strong>of</strong> <strong>the</strong> six <strong>British</strong> species <strong>of</strong> Odon<strong>to</strong>myia (figs 139- 140) is given<br />
by Brindle (1964a).<br />
Hermetiinae. Although not included in <strong>the</strong> <strong>British</strong> Check List in <strong>the</strong> Handbook series,<br />
<strong>the</strong> larvae <strong>of</strong> Hermetia illucens (L.) (fig. 130) frequently come before en<strong>to</strong>mologists who<br />
deal with economic enquiries involving food stuffs imported in<strong>to</strong> Britain (see Medical<br />
and Veterinary Section and Smith & Chainey, 1989).<br />
Xylomyidae<br />
(Figs: larvae 141-143, pupa 1131)<br />
Regarded until very recently as a subfamily <strong>of</strong> Stratiomyidae, this small but widely<br />
distributed family contains less than I 00 species in 53 genera. Only one genus, Xylomya<br />
( = Solva) is represented in Britain, by 3 species.<br />
The larvae (fig. 143) resemble Stratiomyidae with a shagreened cuticle but are<br />
distinguishable at once by <strong>the</strong> shiny smooth areas on <strong>the</strong> thorax and <strong>the</strong> <strong>to</strong>o<strong>the</strong>d anus<br />
(figs 141-143). They are predaceous or saprophagous and live under <strong>the</strong> bark <strong>of</strong> trees<br />
(oak, poplar, elm, walnut) and in rotting logs.<br />
Brindle (196Ic) keys <strong>the</strong> <strong>British</strong> species.<br />
Xylophagidae<br />
(Figs: larva 144, pupa 1132)<br />
This small family (until recently included in <strong>the</strong> Rhagionidae) consists <strong>of</strong> about 30<br />
species in one genus (Xylophagus) and is restricted <strong>to</strong> <strong>the</strong> Holarctic region.<br />
58
Three species <strong>of</strong> Xylophagus occur in Britain, two <strong>of</strong> which are known in <strong>the</strong> larval<br />
<strong>stages</strong> and keyed by Brindle (1961 b, 1966); <strong>the</strong>y occur in dead wood. X. cinctus DeGeer<br />
(fig. 144) is restricted <strong>to</strong> pine trees but X . ater Meigen is found in deciduous trees<br />
including birch, oak, alder, beech and aspen. The larva <strong>of</strong> X.junki Szih\.dy is unknown<br />
but may well be found in conifers since <strong>the</strong> only known <strong>British</strong> adult was found in<br />
Glenmore Forest, near Aviemore, Scotland.<br />
There has been some controversy over <strong>the</strong> feeding habits <strong>of</strong> larval Xylophagus but<br />
<strong>the</strong>y are certainly carnivorous, probably on beetle larvae. Downes (1953) records<br />
X. ater attacking a small staphylinid larva.<br />
Rhagionidae<br />
(Figs: larvae 145- 151, egg 993, pupa 1133)<br />
The Rhagionidae has a world-wide distribution with 500 species contained in some<br />
20 genera. Four subfamilies are recognised <strong>of</strong> which two occur in Britain. No key <strong>to</strong><br />
subfamilies is given since <strong>the</strong> larvae <strong>of</strong> some genera are unknown, but Brindle (1959)<br />
provides a partial key including those genera now removed <strong>to</strong> o<strong>the</strong>r families.<br />
The larvae <strong>of</strong> Rhagio (fig. 145) and Chrysopilus (fig. 150) are carnivorous but are also<br />
said <strong>to</strong> feed on decaying vegetable material. However, from a study <strong>of</strong> <strong>the</strong> larval<br />
mouthparts, Roberts (1969) concludes that Rhagio is carnivorous, feeding on earthworms<br />
(Oiigochaeta) and s<strong>of</strong>t bodied insect larvae. Brindle (1962g) keys <strong>the</strong> larvae <strong>of</strong><br />
four <strong>of</strong> <strong>the</strong> six <strong>British</strong> species <strong>of</strong> Rhagio. Hobby & Smith (1962a) describe <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong> <strong>of</strong> Chrysopilus cristatus (F.), which differs from Rhagio (fig. 148) in <strong>the</strong> structure<br />
<strong>of</strong> <strong>the</strong> head capsule (figs 149-150). Larvae <strong>of</strong> both genera are found in leaf mould and<br />
moss and o<strong>the</strong>r woodland debris.<br />
Ptiolina larvae feed on mosses on trees and s<strong>to</strong>nes. They are shining green in life and<br />
<strong>the</strong> anal segment has two fleshy lobes, one dorsal, one ventral (figs 147, 151 ).<br />
The larva <strong>of</strong> Spania has not been described but is said <strong>to</strong> be small and <strong>to</strong> mine <strong>the</strong><br />
leaves <strong>of</strong>liverworts (Pellia, etc.).<br />
Symphoromyia larvae occur in damp soil. Some Alaskan species have been described<br />
by Sommerman (1962) and were found <strong>to</strong> feed on <strong>the</strong> larvae <strong>of</strong> o<strong>the</strong>r soil-dwelling<br />
Diptera.<br />
A<strong>the</strong>rix and Atrichops <strong>of</strong> <strong>the</strong> Diptera Check List in <strong>the</strong> Handbooks series are treated<br />
here under A<strong>the</strong>ricidae.<br />
A<strong>the</strong>ricidae<br />
(Figs: larvae 152- 156, eggs 994-995, pupa 1134)<br />
Formerly included as a subfamily in <strong>the</strong> Rhagionidae this family contains some 80<br />
world species in about 6 genera. Two genera occur in Britain, both aquatic in <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong>.<br />
The distinctive larvae <strong>of</strong> A<strong>the</strong>rix are found beneath and amongst s<strong>to</strong>nes on river<br />
beds. When fully grown <strong>the</strong>y leave <strong>the</strong> water and burrow in<strong>to</strong> light gravelly soil under<br />
moss on <strong>the</strong> river bank and pupate. The two <strong>British</strong> species are easily distinguished by<br />
<strong>the</strong> difference in <strong>the</strong> lengths <strong>of</strong> <strong>the</strong> lateral processes on <strong>the</strong> abdomen (figs 154-155).<br />
A<strong>the</strong>rix ibis (F.) is well known for <strong>the</strong> curious, and frequently described oviposition<br />
habits <strong>of</strong> <strong>the</strong> female. The flies ga<strong>the</strong>r in clusters on twigs and branches overhanging <strong>the</strong><br />
water and remain <strong>the</strong>re until <strong>the</strong>y die in a mass <strong>of</strong> flies and eggs (fig. 994). The eggs (fig.<br />
995) ultimately hatch and <strong>the</strong> young larvae fall in<strong>to</strong> <strong>the</strong> water. Mackey & Brown (1980)<br />
describe <strong>the</strong> pupa and provide a key <strong>to</strong> pupae for <strong>the</strong> families <strong>of</strong> aquatic Brachycera.<br />
Brindle (1961d) keys <strong>the</strong> species <strong>of</strong> A<strong>the</strong>rix.<br />
59
The larva <strong>of</strong> Atrichops crassipes (Meigen), <strong>the</strong> only o<strong>the</strong>r <strong>British</strong> representative <strong>of</strong><br />
this family, remains undescribed, but its life his<strong>to</strong>ry will probably prove similar <strong>to</strong><br />
that <strong>of</strong> A<strong>the</strong>rix. Adults used <strong>to</strong> occur (1900-1904) on alders in a water-meadow near<br />
Ticehurst Road Station, Sussex, and near Milford-on-Sea, Hants, as well as a few<br />
localities in <strong>the</strong> New Forest (Hants). It has recently been rediscovered in some <strong>of</strong>t he old<br />
localities, and Gibbs (1987) records it from <strong>the</strong> Kennet Valley in Berkshire.<br />
Tabanidae<br />
(Figs: larvae 157- 160, eggs 996-997, pupa 1135)<br />
The large family Tabanidae (horse-flies, clegs, deer flies) contains some 3500 species<br />
and is world-wide in distribution. Adult female Tabanidae suck blood and some are <strong>of</strong><br />
veterinary and medical importance.<br />
The larvae <strong>of</strong> Tabanidae are found in wet mud at <strong>the</strong> margins <strong>of</strong> ponds, lakes and<br />
streams, in sand on <strong>the</strong> sea shore or in tree-holes. The first two instars do not feed and<br />
moulting from first <strong>to</strong> second in star <strong>of</strong>ten takes place on <strong>the</strong> egg mass. The later ins tars<br />
are very active preda<strong>to</strong>rs on worms, molluscs and larvae <strong>of</strong> o<strong>the</strong>r Diptera (including<br />
<strong>the</strong>ir own kind) but <strong>the</strong>y do not feed on Crustacea or beetle larvae. The number <strong>of</strong><br />
larval instars (fig. 157) ranges from 7 <strong>to</strong> 11 and varies, along with <strong>the</strong> time <strong>of</strong> larval<br />
development, even in <strong>the</strong> same species, according <strong>to</strong> a number <strong>of</strong> fac<strong>to</strong>rs, e.g. food,<br />
temperature, humidity.<br />
The larvae can survive for a long time without food (up <strong>to</strong> several months) and in<br />
unfavourable environmental conditions <strong>the</strong>y can hibernate/aestivate several times,<br />
with additional larval instars.<br />
Tabanid larvae can be roughly divided in<strong>to</strong> three ecological groups:<br />
I) Hydrophilous - developing only in water (e.g. Chrysops, some Hybomitra).<br />
2) Semihydrophilous- with first and second instars on surface <strong>of</strong> water or moist soil and in<br />
third instar migrate in<strong>to</strong> soil close <strong>to</strong> water (e.g. most European Tabanidae).<br />
3) Edaphic - living in drier soil, usually far from water (e.g. some HOI!nw<strong>to</strong>pota).<br />
Eggs (figs 996-997) are laid in masses which may consist <strong>of</strong> a single layer (some<br />
Chrysops), two or three layers (Haema<strong>to</strong>pota) or three or four layers (o<strong>the</strong>r tabanids).<br />
The number <strong>of</strong> eggs laid ranges from 400 in small species <strong>to</strong> <strong>10</strong>00 in <strong>the</strong> larger species.<br />
There are 28 <strong>British</strong> species representing 5 genera. Two sub-families and three tribes<br />
occur in Britain, which may be distinguished in <strong>the</strong> mature larval <strong>stages</strong> as follows.<br />
Four tubular pseudopods (ventro-lateral (vlp) and lateral pairs (lp)) on each abdominal<br />
segment except <strong>the</strong> last (fig. 159) . . . . . . . . . Ckysopsinae (Chrysopsini)<br />
At least six tubular pseudopods (2 pairs ventro-lateral, plus a lateral pair) on all but <strong>the</strong> last<br />
abdominal segment (fig. 158) . . . . . . . . . . . . . Tabaninae 2<br />
2 <strong>An</strong>al segment shorter than its own vertical height (fig. 157) . . . . Haema<strong>to</strong>potini<br />
<strong>An</strong>al segment elongated, longer than its own vertical height (fig. 160)<br />
Tabanini<br />
The European Tabanidae have been monographed by Chvala et al. (1972) who<br />
include much biological information. O<strong>the</strong>r important papers on larval Tabanidae are<br />
listed by Jezek (1977), who keys <strong>the</strong> last instar larvae and pupae <strong>of</strong> many species.<br />
Chrysopsinae. Only one genus, Chrysops, occurs in Britain with four species. The<br />
larvae are usually found in mud or sand at <strong>the</strong> edges <strong>of</strong> streams and brooks and 3 <strong>of</strong> <strong>the</strong><br />
4 <strong>British</strong> species are described by Jezek (1970). The larva <strong>of</strong> C. sepu/cralis (F.) is<br />
unknown but should be sought in swampy conditions, especially peat bogs, where <strong>the</strong><br />
adults have a very local distribution.<br />
60
Tabaninae<br />
Haema<strong>to</strong>potini. Only one genus, Haema<strong>to</strong>pota, with 4 species occurs in Britain.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> commonest species, H. pluvialis (L.) (<strong>the</strong> cleg or dun fly)<br />
have been studied in detail by Cameron (1934). The larvae <strong>of</strong> H. crassicornis Wahlberg<br />
also occur in marshy soil but <strong>the</strong> species is commoner than H. pluvialis in upland areas<br />
in <strong>the</strong> hilly districts <strong>of</strong> <strong>the</strong> north <strong>of</strong> England and Scotland. H. grandis Meigen and H .<br />
bigoti Gobert are salt marsh species in Britain but <strong>the</strong> larvae are undescribed.<br />
Brindle (196la) keys <strong>the</strong> larvae <strong>of</strong> three <strong>British</strong> species, including H. italica Meigen<br />
from salt marshes. H. italica however, included only as a dubious synonym <strong>of</strong> H.<br />
grand is in <strong>the</strong> <strong>British</strong> Check List, is regarded as a subspecies <strong>of</strong> H. grandis by Oldroyd in<br />
his key <strong>to</strong> adults in <strong>the</strong> Handbook series, and as a distinct species with larval habits<br />
resembling those <strong>of</strong> H. pluvialis by Chvala et al. (1972) (and <strong>the</strong> larva <strong>of</strong> H . grandis<br />
stated <strong>to</strong> be unknown). Clearly <strong>the</strong> genus Haema<strong>to</strong>pota needs fur<strong>the</strong>r attention by<br />
<strong>British</strong> Dipterists, accompanied by careful rearing from larvae but no genuine <strong>British</strong><br />
adults <strong>of</strong> H. italica are known.<br />
Tabanini. Only two <strong>of</strong> <strong>the</strong> 13 European ( 4 <strong>British</strong>) species <strong>of</strong> Atylotus are known in<br />
<strong>the</strong> larval stage. A .fulvus (Meigen) has been reared from <strong>the</strong> periphery <strong>of</strong> forest ponds<br />
in Czechoslovakia.<br />
Jdek (1977) keys <strong>the</strong> last ins tar larvae and pupae <strong>of</strong> 8 <strong>of</strong> <strong>the</strong> 37 described European<br />
species <strong>of</strong> Hybomitra, including 4 <strong>of</strong> <strong>the</strong> 9 <strong>British</strong> species. H. distinguenda (Verrall) is a<br />
widespread and <strong>of</strong>ten numerous species whose larvae have been found in living or dead<br />
vegetation at <strong>the</strong> margins <strong>of</strong> marshes, forest ponds, marshy meadows, margins <strong>of</strong><br />
drainage near cattle pens and in a moist rotten pine stump. H . bimaculata (Macquart),<br />
H . ciureai (Seguy) (accidentally indented as a synonym in <strong>the</strong> Check List) and H.<br />
muehlfeldi (Brauer) have all been found at <strong>the</strong> edges <strong>of</strong> ponds, usually submerged even<br />
during <strong>the</strong> winter months among frozen Phragmites, Carex and Juncus.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> only 3 <strong>of</strong> <strong>the</strong> 8 <strong>British</strong> species <strong>of</strong> Tabanus have been<br />
adequately described. T. autumnalis L. has been found in a wide variety <strong>of</strong> substrates<br />
from <strong>to</strong>tally organic soil in Alnus forest <strong>to</strong> <strong>the</strong> margins <strong>of</strong> forest ponds and drainage<br />
ditches. T. bromius L. occurs in <strong>the</strong> periphery <strong>of</strong> forest ponds and in molehills; T.<br />
maculicornis Zetterstedt has also been recorded from molehills, but usually in <strong>the</strong><br />
margins <strong>of</strong> peat bogs and in forest meadows near lakes and streams; T. miki Brauer<br />
occurs in <strong>the</strong> mud <strong>of</strong> swamps and marshy meadows (with Phragmites, Carex, Iris and<br />
Salix ).<br />
Most <strong>of</strong> <strong>the</strong> ecological information given above comes from Czechoslovakian data<br />
included in <strong>the</strong> references cited in Jezek ( 1977). It is strange that <strong>the</strong> life-his<strong>to</strong>ries <strong>of</strong> this<br />
important family have been so neglected by <strong>British</strong> en<strong>to</strong>mologists and clearly <strong>the</strong>re is<br />
scope for a substantial research effort here.<br />
Asilidae<br />
(Figs: larvae 165- 176, eggs 998- 999, pupa 1136)<br />
About 5000 species <strong>of</strong> Asilidae (robber flies) in over 400 genera and subgenera have<br />
been described throughout <strong>the</strong> world, <strong>of</strong> which 27 species in 15 genera occur in Britain.<br />
The long white larvae prefer drier habitats, e.g. sandy soil, and are predaceous,<br />
although <strong>the</strong> Laphriinae, which occur in decayed wood, are considered <strong>to</strong> be saprophagous<br />
or herbivorous. A classic work on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> is that <strong>of</strong> Melin<br />
(1923); Brindle (1962e) keys <strong>the</strong> known larvae <strong>of</strong> <strong>the</strong> <strong>British</strong> species. A useful world<br />
review <strong>of</strong> <strong>the</strong> known biology <strong>of</strong> Asilidae is provided by Knutson (1972). Many larvae<br />
are predaceous on beetle larvae but hardly any observations on larval food have been<br />
made for <strong>the</strong> genera occurring in Britain.<br />
61
The higher classification <strong>of</strong> <strong>the</strong> adults <strong>of</strong> Asilidae based on adult structures appears<br />
<strong>to</strong> be very subjective, each specialist creating a different system, e.g. Lep<strong>to</strong>gaster varies<br />
in status from tribe <strong>to</strong> separate family! For <strong>the</strong> purpose <strong>of</strong> this Handbook I follow<br />
Brindle (1962e) in recognising 4 subfamilies for <strong>the</strong> separation <strong>of</strong> <strong>the</strong> larval <strong>stages</strong> as<br />
follows.<br />
Mandibles absent; maxillae (mx) widely separated distally being prevented from closer<br />
approximation by <strong>the</strong> strongly broadened basal part; outer margin <strong>of</strong><strong>the</strong> maxillae with an<br />
incision in which lie <strong>the</strong> maxillary palps (mp) (figs 165-166) . . . . Lep<strong>to</strong>gastrinae<br />
Mandibles (md) present; maxillae usually closely approximated along <strong>the</strong>ir length, basal<br />
part not broadened internally . . . . . . . . . . . . . . . . . . 2<br />
2 Maxillae narrow, without a lateral incision (fig. 167); anal segment '"'ith a horizontal apical<br />
brown or white ridge (fig. 169, r) . . . . . . . . . . . . . . . Asilinae<br />
Maxillae broad, with a lateral incision in which lie <strong>the</strong> maxillary palps (fig. 170); anal segment<br />
without an apical ridge . . . . . . . . . . . . . . . . . . . . 3<br />
3 Abdominal segments 1-{i with an annular ring <strong>of</strong> 3 or 4 pairs <strong>of</strong> tubercles (fig. 171); anal<br />
segment with an apical sclerotised plate (figs 171 - 173) . . . . . . . Laphriinae<br />
Abdominal segments 1-{i with only one pair <strong>of</strong> ventral tubercles; anal segment without an<br />
apical sclerotised plate (figs 174-176) . . . . . . . Dasypogoninae<br />
Lep<strong>to</strong>gastrinae. There is only one <strong>British</strong> genus, Lep<strong>to</strong>gaster, with two species, <strong>the</strong><br />
larvae <strong>of</strong> which occur in sandy soil or meadows with dense ground vegetation (figs<br />
165- 166).<br />
Dasypogoninae. The genera Leptarthrus ( = /sopogon) and Dasypogon ( = Selidopogon),<br />
each with one species, are unknown in <strong>the</strong> larval <strong>stages</strong> though L. brevirostris<br />
(Meigen) has been seen <strong>to</strong> oviposit on cocksfoot and <strong>the</strong> larvae should probably be<br />
sought in soil on chalk downland. Adults <strong>of</strong> D. diadema F. have not been found in<br />
Britain for well over a century but should <strong>the</strong> species still survi,·e in Britain its larvae<br />
should be sought on <strong>the</strong> dunes <strong>of</strong> <strong>the</strong> Welsh coast.<br />
Of <strong>the</strong> six <strong>British</strong> species <strong>of</strong> Dioctria only D. rufipes (De Geer) (figs 174-176) and D.<br />
oelandica (L.) are known in <strong>the</strong> larval stage and <strong>the</strong>y occur in woodland soil (Brindle<br />
1968, 1969a). The larva <strong>of</strong> Lasiopogon cinctus (F.) is not uncommon in sandy soils in<br />
pastures, woodlands and dunes, but also occurs in richer soils.<br />
Laphriinae. The larvae (fig. 171) <strong>of</strong> this subfamily are found in <strong>the</strong> stumps <strong>of</strong> decayed<br />
trees. Laphriaflava L. (figs 172-173) is found in decayed wood <strong>of</strong> pine, spruce or birch<br />
stumps in Scotland. L. gilva (L.) is found locally in sou<strong>the</strong>rn England between <strong>the</strong> bark<br />
and wood <strong>of</strong> pine stumps. L. marginata (L.) is also restricted <strong>to</strong> sou<strong>the</strong>rn England<br />
where it occurs in pine stumps or decayed deciduous wood.<br />
Asilinae. All nine genera <strong>of</strong> <strong>the</strong> subfamily occurring in Britain are lrnown in <strong>the</strong> larval<br />
stage, though not all are known from <strong>British</strong> species or material.<br />
Larvae <strong>of</strong> Asilus crabroniformis (L.) are locally common in sou<strong>the</strong>rn England in <strong>the</strong><br />
sandy soils <strong>of</strong> pastures and heaths. One <strong>of</strong> <strong>the</strong> three species <strong>of</strong> Epitriptus (regarded as<br />
synonymous with Machimus by some workers), E. cingulatus(F.), is lrnown in <strong>the</strong> larval<br />
stage and occurs in <strong>the</strong> sandy soils <strong>of</strong> meadows, grassy slopes, etc. Eu<strong>to</strong>lmus rufibarbis<br />
(Meigen) is found in similar situations. The larvae <strong>of</strong> Machimvs atricapillus (Fallen)<br />
and Neoitamus cyanurus (Loew) occur in sandy soils in open woodlands and <strong>the</strong><br />
unknown larvae <strong>of</strong> <strong>the</strong> o<strong>the</strong>r rarer single species <strong>of</strong> each <strong>of</strong> <strong>the</strong>se two genera will no<br />
doubt be found in similar situations. The larvae <strong>of</strong> Pamponerus germanicus (L.) (fig.<br />
168) and Philonicus a/biceps (Meigen) are found in sand dune soils and Rhadiurgus<br />
variabilis (Zetterstedt) occurs locally in sandy soils at <strong>the</strong>edges<strong>of</strong>pine woods in upland<br />
districts, chiefly in Scotland. The larva <strong>of</strong> Dysmachus trigonus (Meigen) is unknown but<br />
Melin (1923) found that <strong>of</strong> a Swedish species in sandy soils on <strong>the</strong> coast and inland<br />
62
heaths. In <strong>the</strong> <strong>British</strong> Museum (Nat. Hist.) collections <strong>the</strong>re are two pupae <strong>of</strong> D.<br />
trigonus from St. Merryn, Cornwall, and Croyd, N . Devon, one labelled 'in sand' (coli.<br />
K. G. Blair ex coli L. Parmenter).<br />
Therevidae<br />
(Figs: larva 161-162, pupa 1140)<br />
About 700 species <strong>of</strong> Therevidae (stilet<strong>to</strong> flies) have been described throughout <strong>the</strong><br />
world, <strong>of</strong> which 13 species in 3 genera occur in Britain.<br />
The life his<strong>to</strong>ries are very poorly known but <strong>the</strong> vermiform (worm- or snake-like)<br />
larvae (fig. 161 ), with abdominal segments 1-6 subdivided, occur in sandy <strong>to</strong> sandyloam<br />
soils and are preda<strong>to</strong>rs on a variety <strong>of</strong> arthropods and earthworms, with a<br />
preference (like Asilidae) for beetle larvae, especially <strong>of</strong> <strong>the</strong> families Elateridae,<br />
Scarabaeidae and Tenebrionidae. There are five larval <strong>stages</strong> in <strong>the</strong> life-his<strong>to</strong>ries so far<br />
studied.<br />
Thereva is <strong>the</strong> genus most frequently encountered in <strong>the</strong> larval stage (figs 161 - 162).<br />
Fox Wilson (1924) conducted experiments on <strong>the</strong> feeding habits <strong>of</strong> Thereva plebeia (L.)<br />
and concluded that larvae fed on leaf-mould and humus in <strong>the</strong> soil and when this was<br />
scarce would attack living plants, e.g. cabbage and pota<strong>to</strong> and o<strong>the</strong>r garden vegetables.<br />
However, it may well be that this only occurs in <strong>the</strong> absence <strong>of</strong> suitable invertebrate<br />
food, as <strong>the</strong> larvae are undoubtedly voracious preda<strong>to</strong>rs and feed on various insect<br />
larvae including wireworms (Coleoptera, Elateridae) and <strong>the</strong>ir own kind. They are also<br />
capable <strong>of</strong> biting man, usually in sandy areas; Smith (1979a, 1986a) summarises <strong>the</strong>se<br />
records and one more can be added from a dry rocky limes<strong>to</strong>ne area on Bunster Hill,<br />
Dovedale (K.N.A. Alexander, pers. comm.).<br />
Collinge ( 1909) (overlooked by Fox Wilson) made observations on Thereva<br />
nobilitata F. from larvae sent in by gardeners and nurserymen from among <strong>the</strong> roots <strong>of</strong><br />
various plants. He fed <strong>the</strong> larvae on Diptera larvae and small earthworms but noted<br />
that <strong>the</strong>y preferred weevil larvae which were less active. The larvae <strong>of</strong> <strong>the</strong> remaining 8<br />
species <strong>of</strong> Thereva are unknown but should be sought in sandy soil; T. annulata F. is<br />
common on coastal sand dunes.<br />
Larvae <strong>of</strong> Psilocephala melaleuca (Loew) have been found in rotting stumps in <strong>the</strong><br />
Windsor Forest- Ascot area <strong>of</strong> Berkshire, but have not been described.<br />
The larva <strong>of</strong> Dialineura ani/is (L.) is unknown, but should be sought on sandhills <strong>of</strong><br />
<strong>the</strong> Welsh and Scottish coasts where adults are common.<br />
Scenopinidae<br />
(Figs: larva 163-164, egg I 000, pupa 1138)<br />
Some 312 species <strong>of</strong> Scenopinidae (window flies, carpet flies) have been described<br />
throughout <strong>the</strong> world. The family is represented in Britain by 3 species in <strong>the</strong> single<br />
genus Scenopinus.<br />
The slender snake-like larvae resemble those <strong>of</strong> Therevidae in having abdominal<br />
segments 1- 6 subdivided (see key) and are predaceous on <strong>the</strong> larvae and pupae <strong>of</strong><br />
clo<strong>the</strong>s-moths, beetles and fleas . They are frequently found in carpets but since this can<br />
hardly be <strong>the</strong>ir 'natural' habitat it is assumed that <strong>the</strong> debris in birds' nests provided<br />
<strong>the</strong>ir original larval pabulum. They have been recorded most frequently from nests <strong>of</strong><br />
house sparrows, swallows, pigeons, starlings and jackdaws (Hicks, 1959). Scenopinid<br />
larvae are no<strong>to</strong>riously resistant <strong>to</strong> desiccation and lack <strong>of</strong> oxygen. There is one record<br />
<strong>of</strong> involvement in human urogenital myiasis (Thompson, J.H. et al., 1970).<br />
Two <strong>of</strong> <strong>the</strong> three <strong>British</strong> species, S. fenestra/is (L.) and S. glabrifrons Meigen, are<br />
world wide in distribution but only <strong>the</strong> larva <strong>of</strong> S. fenestra/is (fig. 164) has been<br />
63
described in any detail (Seguy, 1921), while Zuranska (1979) has described <strong>the</strong><br />
puparium.<br />
Acroceridae<br />
(Figs: larvae 177-182, eggs I 00 I, pupa 1139)<br />
Some 475 species in 48 genera <strong>of</strong> Acroceridae have been described throughout <strong>the</strong><br />
world. All are internal parasites <strong>of</strong> spiders (Araneae), particularly Lycosidae, but also<br />
<strong>of</strong> Theridiidae and Gnaphosidae. The black eggs (fig. <strong>10</strong>0 I) may be laid in flight and<br />
scattered on <strong>the</strong> ground in large numbers or deposited in masses on grass stems,<br />
horsetails, twigs or tree trunks away from <strong>the</strong> intended hosts. The first instar larva (figs<br />
179-181) is an active 'planidium' which seeks out and burrows in<strong>to</strong> <strong>the</strong> host. These<br />
young larvae can jump for distances <strong>of</strong> 5 or 6 mm by bending and suddenly straightening<br />
<strong>the</strong> body, using caudal processes and a sucker attachment. They can also move<br />
along a spider's thread with a movement resembling that <strong>of</strong> a loo per caterpillar. The<br />
larvae <strong>of</strong> some species are known <strong>to</strong> obtain <strong>the</strong>ir oxygen by applying <strong>the</strong>ir posterior<br />
spiracles <strong>to</strong> <strong>the</strong> spider's lung-books. There are three larval instars and <strong>the</strong> mature larva<br />
(fig. 177) usually emerges posteriorly from <strong>the</strong> spider's epigastric furrow area and<br />
pupates free <strong>of</strong> <strong>the</strong> host, <strong>of</strong>ten in a protective webbing spun by <strong>the</strong> host spider before its<br />
death.<br />
Only three species (Acrocera globulus Panzer, Ogcodes gibbosus L. and 0. pallipes<br />
Latreille) occur in Britain and <strong>the</strong>se are very rarely seen even as adults. They occur in<br />
scattered localities from Herefordshire southwards and may be locally abundant.<br />
Since spiders are so common it is strange that Acroceridae are so rarely encountered.<br />
Spiders are known <strong>to</strong> kill some <strong>of</strong> <strong>the</strong> larvae and <strong>the</strong> adult flies are frequently caught in<br />
<strong>the</strong>ir webs but in view <strong>of</strong> <strong>the</strong> large numbers <strong>of</strong> eggs laid (80Q-900 have been observed)<br />
<strong>the</strong> mortality seems high and a quantitative investigation would be worth while for<br />
anyone with access <strong>to</strong> a local colony.<br />
Bombyliidae<br />
(Figs: larvae 183-188, egg I 002, pupa 1137)<br />
Nearly 4000 species <strong>of</strong> Bombyliidae (bee-flies) in some 194 genera have been<br />
described throughout <strong>the</strong> world. The family is best represented in arid and semi-arid<br />
regions. Few species are known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> but all are parasitic on <strong>the</strong> larvae<br />
or pupae (or both) <strong>of</strong> various species <strong>of</strong> Lepidoptera, Hymenoptera, Coleoptera,<br />
Diptera and Neuroptera, or predaceous on <strong>the</strong> egg pods <strong>of</strong> grasshoppers. There are 3<br />
larval <strong>stages</strong>, <strong>the</strong> first being a planidium (fig. 183). As with Asilidae, <strong>the</strong> pupal stage (fig.<br />
1137) is better described because so <strong>of</strong>ten only <strong>the</strong> pupal skin is recovered after a chance<br />
rearing. Hull (1973) provides generic keys <strong>to</strong> <strong>the</strong> pupal stage. Eggs (fig. <strong>10</strong>02) are said <strong>to</strong><br />
be produced in large numbers. Du Merle (1975) provides a host list for <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong>.<br />
Only 11 species in 5 genera occur in Britain. The higher classification <strong>of</strong> <strong>the</strong><br />
Bombyliidae is ra<strong>the</strong>r unstable. Of <strong>the</strong> 12 or so subfamiles recognised 4 are represented<br />
in Britain but <strong>the</strong> larvae are <strong>to</strong>o imperfectly known <strong>to</strong> construct a key.<br />
<strong>An</strong>thracinae. The only <strong>British</strong> member <strong>of</strong> this subfamily, <strong>the</strong> very rare (Leicestershire<br />
only) <strong>An</strong>thrax anthrax (Schrank), has not been described in <strong>the</strong> larval stage, though <strong>the</strong><br />
pupa is known. Abroad it has been reared from <strong>the</strong> nest <strong>of</strong> Odynerus spinipes (L.)<br />
(Hym., Vespidae); from both Osmia nigriventris (Zetterstedt) and its parasite Stelis<br />
ornatula Klug (both Hym., Apidae); and from nests <strong>of</strong> <strong>An</strong>thophora and Megachile<br />
(both Hym., Apidae).<br />
64
Bombyliinae. T. A. Chapman (1878) found pupae <strong>of</strong> Bombylius major L., <strong>the</strong><br />
commonest <strong>British</strong> species, in <strong>the</strong> nest <strong>of</strong> <strong>the</strong> mining bee <strong>An</strong>drena labia/is Kirby. O<strong>the</strong>r<br />
bees have been recorded as hosts on <strong>the</strong> continent, e.g. A. humilis Imh<strong>of</strong>f and Colletes<br />
cunicularia (L.). T. A. Chapman also observed females <strong>of</strong> a small brown species <strong>of</strong><br />
Bombylius (probably canescens Mikan) laying its eggs in flight over a bank containing<br />
Ha/ictus burrows. On <strong>the</strong> continent <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> B. minor L. have been<br />
described from nests <strong>of</strong> Colletes daviesana Smith (Nielsen, J. C., 1903). The first stage<br />
larvae <strong>of</strong> B. minor feed on <strong>the</strong> pollen and honey s<strong>to</strong>re but in later instars attack <strong>the</strong> bee<br />
larvae.<br />
Exoprosopinae. The larva (fig. 186) <strong>of</strong> Thyridanthrax fenestratus (Fallen) has been<br />
described by Seguy (1932). It has been recorded from <strong>the</strong> egg pods <strong>of</strong> grasshoppers in<br />
<strong>the</strong> Mediterranean area but Grea<strong>the</strong>ad (1963) has suggested that in at least some <strong>of</strong><br />
<strong>the</strong>se cases it may be parasitic on Diptera associated with <strong>the</strong> egg pods. The species has<br />
not been reared in Britain but adults occur locally on sandy commons in sou<strong>the</strong>rn<br />
England. Grasshoppers and associated Diptera in such localities could well repay<br />
investigation.<br />
Larvae <strong>of</strong> Villa species parasitize larvae <strong>of</strong> Lepidoptera (especially Noctuidae) and<br />
Coleoptera (Tenebrionidae). Of <strong>the</strong> three <strong>British</strong> species, only V. circumdata (Meigen)<br />
has been reared from a pyralid moth larva (in France).<br />
Phthiriinae. Phthiria pulicaria Mikan is a sand-dune species and has been reared from<br />
Scrobipalpula psilella (Herrich-Schiiffer) (Lep., Gelechiidae) on <strong>the</strong> continent.<br />
Empididae<br />
(Figs: larvae 189-205, eggs <strong>10</strong>03- <strong>10</strong>04, pupae 1146-1148)<br />
Over 3000 species <strong>of</strong> Empididae (dance flies) have been described throughout <strong>the</strong><br />
world and many more await description. Over 350 species have so far been recorded in<br />
Britain. The family has usually been divided in<strong>to</strong> 8 or <strong>10</strong> subfamilies <strong>of</strong> which 6 occur in<br />
Britain. Chvala ( 1983) has grouped <strong>the</strong>se in<strong>to</strong> distinct families: Microphoridae, Atelestidae<br />
(Atelestus is under Platypezidae in <strong>the</strong> Check List, larvae unknown), Hybotidae<br />
(including Ocydromiinae, Hybotinae and Tachydromiinae), and Empididae (including<br />
Oreoge<strong>to</strong>ninae, Empidinae, Brachys<strong>to</strong>matinae (not <strong>British</strong>), Hemerodromiinae and<br />
Clinocerinae). The group is here treated as one family, since <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are so<br />
poorly known and <strong>the</strong> available information is given under <strong>the</strong> six sub-family headings<br />
included in <strong>the</strong> Check List (Smith, 1976) in this Handbook series. The following key<br />
couplet will facilitate preliminary identification <strong>of</strong> larvae.<br />
Keys <strong>to</strong> <strong>the</strong> few genera so far known in <strong>the</strong> larval <strong>stages</strong> are given by Smith (1969) and<br />
McAipine et al. (1981).<br />
Larvae terrestrial, with amphipneustic respira<strong>to</strong>ry system (spiracles fairly conspicuous),<br />
without ventral abdominal pseudopods (figs 196-200) . . . . . . . . . . . .<br />
. . . . . . . . . . . Tachydromiinae, Hybotinae, Ocydromiinae, Empidinae<br />
Larvae aquatic or semi-aquatic, with apneustic respira<strong>to</strong>ry system (no spiracles) and with<br />
ventral abdominal pseudopods (figs 201- 205) Hemerodromiinae, Clinocerinae<br />
Tachydromiinae. There is no adequate description <strong>of</strong> a larva <strong>of</strong> this subfamily. I have<br />
seen Crossopalpus curvipes Meigen reared from a cocoon in a dead snail ( Cepaea<br />
nemoralis L.) by R. A. Beaver and <strong>the</strong> mouth-parts and posterior spiracles, recovered<br />
from <strong>the</strong> cast skin, are illustrated (figs 190-192). Malloch (1917) figured <strong>the</strong> mouth parts<br />
<strong>of</strong> Drapetis assimilis Fallen (as nigra Meigen) which I have included for comparison<br />
(fig. 189). Malloch noted that <strong>the</strong> Drapetis larva occurs under bark or in decaying wood<br />
65
and makes a <strong>to</strong>ugh cocoon in which it pupates. Be ling ( 1888) gives a brief description <strong>of</strong><br />
<strong>the</strong> larva <strong>of</strong> Platypalpus major (Zetterstedt) found in soil under moss in a wood. The<br />
rarity or absence <strong>of</strong> males in some species <strong>of</strong> Platypalpus has led <strong>to</strong> <strong>the</strong> suggestion that<br />
<strong>the</strong>y may be entirely par<strong>the</strong>nogenetic (P. major (Zetterstedt)) or at least so over part <strong>of</strong><br />
<strong>the</strong>ir range (e.g. P. candicans (Fallen) in N. Europe, P. cursitans (F.) in central and<br />
sou<strong>the</strong>rn Europe).<br />
There are some records <strong>of</strong> reared adults which indicate where <strong>the</strong> larvae, which<br />
almost certainly are all predaceous, should be sought, e.g. Crossopalpus, dung,<br />
fungi (C. nigritellus Zetterstedt in young plasmodium <strong>of</strong> Fuligo septica Gmelin<br />
(Myxomycetes)); Elaphropeza ephippiata (Fallen), woodland soil (G. C. Varley);<br />
Tachypeza nubila (Meigen), under bark, in fungi (e.g. Bjerkandera, Amanita, Paxillus,<br />
Hypholoma, Daldinia).<br />
The remaining genera should be sought where adults are known <strong>to</strong> occur, especially<br />
in soil and decaying wood, or soil under grass-tufts (Stilpon) sand on <strong>the</strong> sea shore,<br />
rivers or lakes (Chersodromia, some Tachydromia).<br />
Hybotinae. The <strong>immature</strong> <strong>stages</strong> are unknown but <strong>the</strong> larvae <strong>of</strong> Hybos probably live<br />
in soil around woodland margins or scrubland. The larvae <strong>of</strong> Syndyas should be sought<br />
in cold Sphagnum bogs. Syneches muscarius (F.), <strong>the</strong> only <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus,<br />
should be sought in wet meadows though only one locality is yet known in Britain (The<br />
Moors, Wool, Dorset).<br />
Ocydromiinae. Ocydromia glabricula (Fallen) females are viviparous and scatter<br />
young larvae while in flight over dung, in which <strong>the</strong>y develop (Hobby & Smith, 1962c).<br />
It is probable that <strong>the</strong> o<strong>the</strong>r <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus, 0. melanopleura Loew, is<br />
viviparous as Chvala ( 1983) has seen small dead larvae attached <strong>to</strong> <strong>the</strong> tip <strong>of</strong> <strong>the</strong><br />
abdomen <strong>of</strong> dried female specimens.<br />
Bicellaria, Trichina and Trichinomyia occur in moist shady places, in deciduous<br />
woodland and <strong>the</strong>ir larvae are probably in <strong>the</strong> soil. Euthyneura, Oedalea and Lep<strong>to</strong>peza<br />
have all been reared from rotten wood and probably Oropezella occurs <strong>the</strong>re <strong>to</strong>o, but<br />
<strong>the</strong> larvae <strong>of</strong> all remain undescribed.<br />
Empidinae. The larvae <strong>of</strong> Empis (fig. 196) and Rhamphomyia (fig. 198) are preda<strong>to</strong>rs,<br />
probably mostly on o<strong>the</strong>r Diptera larvae in woodland soil under leaf-litter (Hobby<br />
& Smith, 1961a, b, 1962b). Larvae <strong>of</strong> Rh. sulcata Meigen may occur in large numbers<br />
in freshly ploughed pasture soil (Smith, 1968). On <strong>the</strong> continent Rh. dentipes<br />
Zetterstedt has been reared from a beech stump and Dr B. R. Laurence has reared Rh.<br />
crassirostris (Fallen) from a pupa found under moss and liverworts on a tree stump.<br />
The larvae <strong>of</strong> Hilara may be distinguished from Empis and Rhamphomyia by <strong>the</strong><br />
larger posterior lobe (fig. 197) which is curved <strong>to</strong>wards <strong>the</strong> tip; <strong>the</strong>y are found in moist<br />
woodland soil under leaf litter, in mole casts and in decaying wood.<br />
Hemerodromiinae. The larvae <strong>of</strong> this subfamily (except Phyllodromia (see below)) are<br />
aquatic and live in mosses in rivers and streams including hygropetricous habitats e.g.<br />
mosses growing in <strong>the</strong> water flowing down vertical rock faces or on <strong>the</strong> tufa (which <strong>the</strong>y<br />
burrow in<strong>to</strong>) deposited by lime-rich waters. The larvae have 7 pairs <strong>of</strong> retractable<br />
pseudo pods on <strong>the</strong> abdominal segments but <strong>the</strong> arrangement and length <strong>of</strong> processes<br />
on <strong>the</strong> anal segment varies between genera and species (figs 202-205).<br />
Hemerodromia unilineata (Zetterstedt) (fig. 202) larvae occur most commonly in<br />
mosses in deeper water where <strong>the</strong> current is slow. The larvae <strong>of</strong> Chelifera trapezina<br />
(Zetterstedt) are found in very fast flowing water, e.g. where <strong>the</strong> water runs down<br />
sloping artificial weirs. The striking pupae <strong>of</strong> <strong>the</strong>se two genera have <strong>the</strong> spiracles <strong>of</strong><br />
<strong>the</strong> prothorax and first seven abdominal segments modified in<strong>to</strong> long slender gills<br />
(fig. 1146).<br />
66
The larva <strong>of</strong> Phyllodromia is terrestrial and <strong>the</strong> anal segment has a large posterior<br />
lobe resembling Hilara. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> o<strong>the</strong>r <strong>British</strong> genera are unknown.<br />
Brindle (1964c, 1969b) keys <strong>the</strong> <strong>immature</strong> <strong>stages</strong> (including Clinocerinae).<br />
Clinocerinae. The larvae <strong>of</strong> this subfamily, like <strong>the</strong> Hemerodromiinae, are aquatic,<br />
but have 8 pairs (fig. 201) <strong>of</strong> abdominal pseudopods.<br />
Vaillant (1967) has studied <strong>the</strong> ecological requirements <strong>of</strong> Wiedemannia in French<br />
rivers and streams and proposes seven ecological zones. He found that <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong> <strong>of</strong> many species may be distributed along <strong>the</strong> same watercourse, depending upon<br />
<strong>the</strong> elevation, rate <strong>of</strong> flow, abundance <strong>of</strong> o<strong>the</strong>r aquatic fauna, and especially <strong>the</strong> maximal<br />
and minimal temperatures <strong>of</strong> <strong>the</strong> water throughout <strong>the</strong> year. Brindle (1969b) found<br />
larvae <strong>of</strong> W. rhynchops (Nowicki) in rivers and streams with a wide range <strong>of</strong> current<br />
speeds but <strong>the</strong>y were fewer in numbers than Hemerodromiinae. Wiedemannia larvae<br />
have been found <strong>to</strong> feed on Simulium larvae (in Algeria).<br />
The larva <strong>of</strong> C/inocera stagna/is (Haliday) has been found in a small shallow lake,<br />
also among filamen<strong>to</strong>us algae on s<strong>to</strong>nes in a slow moving stream (in company with<br />
Wiedemannia Iota Walker and probably feeding on chironomid larvae (Brindle 1973)).<br />
Hin<strong>to</strong>n (1950) showed that <strong>the</strong> pupa <strong>of</strong> Clinocera is adapted <strong>to</strong> survive both in and<br />
out <strong>of</strong> water, providing a certain level <strong>of</strong> relative humidity obtains. C. bipunctata<br />
(Haliday) pupates in mosses just above <strong>the</strong> water level and if <strong>the</strong> water level rises <strong>the</strong><br />
pupa wriggles up through <strong>the</strong> mosses <strong>to</strong> a position just above <strong>the</strong> new level. Pupae <strong>of</strong><br />
W. rhynchops have been found in similar situations and presumably react in a similar<br />
way. Brindle ( 1969b) suggests that <strong>the</strong> differences in pupation sites <strong>of</strong>Hemerodromiinae<br />
and Clinocerinae are correlated with <strong>the</strong> degree <strong>of</strong> development <strong>of</strong> <strong>the</strong> pupal spiracles<br />
(i.e. long gills in <strong>the</strong> former and <strong>of</strong> normal length in <strong>the</strong> latter).<br />
Brindle ( 1964c, 1969b, 1973) provides keys <strong>to</strong> <strong>the</strong> <strong>the</strong>n known genera and species<br />
<strong>of</strong> <strong>the</strong> subfamily. Vaillant (1952) has described <strong>the</strong> aquatic <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
Dolichocephala ocellata (Costa) (fig. 204) from France and provides a revised key <strong>to</strong><br />
genera for <strong>the</strong> sub-family.<br />
Dolichopodidae<br />
(Figs: larvae 206--223, 226; eggs <strong>10</strong>05- <strong>10</strong>06; cocoons 224, 228, 1142, 1145;<br />
pupae 225, 227, 1141- 1145; stem mine 1291)<br />
Some 4500 species <strong>of</strong> Dolichopodidae have been described throughout <strong>the</strong> world.<br />
The <strong>immature</strong> <strong>stages</strong> are not well known but <strong>the</strong> larvae appear <strong>to</strong> be predaceous and are<br />
found in damp soil, sand, rotting wood, under bark, in sap exudations on trees, and in<br />
cattle dung; some are stem-miners and some are aquatic, including <strong>the</strong> inter-tidal zone<br />
<strong>of</strong> <strong>the</strong> sea shore. Pupation usually takes place in a cocoon <strong>of</strong> sand, mud or a piece <strong>of</strong><br />
wood (figs 224--228, 1142-1145). There are 267 <strong>British</strong> species in 38 genera. The family<br />
is divided in<strong>to</strong> nine subfamilies in Britain but <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are <strong>to</strong>o poorly known<br />
<strong>to</strong> provide a key. Keys <strong>to</strong> genera for <strong>the</strong> larvae so far described are given in McAipine et<br />
al. ( 1981 ). A useful 'spot' character for distinguishing most Dolichopodidae from o<strong>the</strong>r<br />
Brachycera larvae (especially Empididae with which <strong>the</strong>y are <strong>of</strong>ten found), is <strong>the</strong><br />
swollen posterior tip <strong>to</strong> each <strong>of</strong> <strong>the</strong> 'rods' <strong>of</strong> <strong>the</strong> mouthparts (fig. 215). O<strong>the</strong>r larval and<br />
pupal differences are discussed by Dyte ( 1967).<br />
Sciapodinae. Only <strong>the</strong> genus Sciapus occurs in Britain, represented by 8 species, one<br />
<strong>of</strong> which is known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>. Larvae <strong>of</strong> Sciapus platypterus (F.) were<br />
found in soil under leaves in a forest <strong>of</strong> beech trees in April by Beling who gave a brief<br />
description, later translated by Lundbeck (1912).<br />
67
Dolichopodinae. Of <strong>the</strong> 53 species <strong>of</strong> Dolichopus occurring in Britain only 16 are<br />
known at all in <strong>the</strong> larval <strong>stages</strong> and <strong>the</strong>se mostly from continental material. Few are<br />
adequately described. Early rearings by Beling, Brauer and o<strong>the</strong>rs are summarised by<br />
Lundbeck (1912). Dolichopus claviger Stannius, D. discifer Stannius, D. /atelimbatus<br />
Macquart, D.lepidus Staeger, D./ongicornis Stannius, D. popular is Wiedemann and D.<br />
ungulatus (L.) have all been reared from soil in meadows or woods. In addition larvae<br />
and pupae <strong>of</strong> D. popular is have been found under moss on old 'stubs' <strong>of</strong> beeches and D.<br />
ungulatus has been found in hollow poplars. Vaillant (1950) described <strong>the</strong> egg (fig.<br />
<strong>10</strong>06), larva and pupa <strong>of</strong> D. griseipennis Stannius reared from eggs laid in moist soil<br />
and <strong>the</strong> same species has been reared from wet grit (Dyte, 1959) and old cow dung<br />
(Laurence, 1953). Dyte (1959) has reared D. nubilus Meigen from intertidal mud in<br />
estuaries and from <strong>the</strong> muddy bank <strong>of</strong> a freshwater stream and he illustrates <strong>the</strong> pupa<br />
(fig. 1141). Dyte also records rearing D. atratus Meigen, D. pennatus Meigen, D.<br />
signatus Meigen and D. trivia/is Haliday from freshwater mud or moist soil. Nielsen<br />
et al. ( 1954) found larvae <strong>of</strong> D. p/umipes (Scopoli) (fig. 207) in numbers from a range <strong>of</strong><br />
half-dry <strong>to</strong> wet soil in Iceland.<br />
No larvae <strong>of</strong> <strong>the</strong> 22 <strong>British</strong> species <strong>of</strong> Hercos<strong>to</strong>mus have been described but H.<br />
cretifer Walker has been reared from moss in a river bank by B. R. Laurence; H.<br />
nigriplantis Stannius has been reared from debris in <strong>the</strong> fork <strong>of</strong> a tree some 6 feet above<br />
<strong>the</strong> ground (Dyte, 1959) and <strong>the</strong> late Dr B. M. Hobby and myself have reared H.<br />
cupreus Fallen from woodland soil.<br />
Of <strong>the</strong> 4 species <strong>of</strong> Tachytrechus occurring in Britain <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
Tachytrechus notatus (Stannius) have been described by Vaillant (1949) from larvae<br />
found on wet rocks, in company with those <strong>of</strong> Lianca/us virens (Scopoli) in Algeria.<br />
Lundbeck (1912) described <strong>the</strong> pupa <strong>of</strong>T. insignis (Stannius) which was found in a<br />
cocoon made <strong>of</strong> sand grains at <strong>the</strong> border <strong>of</strong> a lake in Denmark.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Hypophyllus and Poeci/obothrus are unknown.<br />
Hydrophorinae. The larvae and cocoons containing <strong>the</strong> pupae <strong>of</strong> Hydrophorus<br />
oceanus (Macquart) (figs 208- 2<strong>10</strong>, 223- 225) occur in intertidal mud in estuaries or in<br />
sand on <strong>the</strong> sea shore (Tsacas, 1959; Dyte, 1959). However, since <strong>the</strong> adults <strong>of</strong> this<br />
species also occur inland <strong>the</strong> <strong>immature</strong> <strong>stages</strong> may be found on <strong>the</strong> s<strong>to</strong>nes <strong>of</strong> freshwater<br />
habitats. Probably larvae <strong>of</strong> <strong>the</strong> o<strong>the</strong>r 8 <strong>British</strong> species will be found <strong>to</strong> occupy similar<br />
habitats.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> See/Ius notatus (F.) are unknown but should probably be<br />
sought at <strong>the</strong> water's edge although <strong>the</strong> adults can be found in wet and dry places.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Liancalus virens are found in mosses on partly submerged<br />
wet rocks, and under waterfalls, etc., and are described by Vaillant (1948). The larva<br />
(fig. 213) is metapneustic and <strong>the</strong> pupa (fig. 1143) is hemipneustic. The prothoracic<br />
spiracles <strong>of</strong> <strong>the</strong> pupa are produced in<strong>to</strong> two very long horns which project through a<br />
narrow opening at <strong>the</strong> anterior end <strong>of</strong> <strong>the</strong> cocoon (fig. 1142). Hin<strong>to</strong>n (1950) found that<br />
<strong>the</strong> pupae, within <strong>the</strong>ir cocoons, can survive prolonged submergence if <strong>the</strong> water is well<br />
aerated and also prolonged periods <strong>of</strong> drying (2 days).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Orthoceratium and Thinophilus are unknown but should be<br />
sought on <strong>the</strong> seashore or salt-marshes.<br />
Aphrosylinae. The one <strong>British</strong> genus, Aphrosylus, contains 4 species normally found<br />
on <strong>the</strong> sea shore. The larvae (fig. 214) have not been adequately described but have<br />
been reared from <strong>the</strong> lit<strong>to</strong>ral region, though some (e.g. A . mitis Verrall) may be<br />
estuarine. They are carnivores and sometimes A. celtiber Haliday lays eggs on<br />
limpets and in <strong>the</strong> shells <strong>of</strong> barnacles. Hin<strong>to</strong>n (1967b) has studied <strong>the</strong> structure <strong>of</strong> <strong>the</strong><br />
respira<strong>to</strong>ry horns <strong>of</strong> <strong>the</strong> pupa <strong>of</strong> Aphrosylus (fig. 1144) in relation <strong>to</strong> those <strong>of</strong> o<strong>the</strong>r<br />
Dolichopodidae.<br />
68
Medeterinae. The larvae <strong>of</strong> Medetera (fig. 216) are found under <strong>the</strong> bark <strong>of</strong> trees and<br />
logs, usually in <strong>the</strong> mines <strong>of</strong> Scolytidae and o<strong>the</strong>r beetles on whose larvae and pupae<br />
<strong>the</strong>y feed. About <strong>10</strong> <strong>of</strong> <strong>the</strong> 27 <strong>British</strong> species have been reared from such situations but<br />
identifications <strong>of</strong> <strong>the</strong> older records are unreliable. R. A. Beaver ( 1966) has described <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> and life his<strong>to</strong>ry <strong>of</strong> M . nitida (Macquart) and M . impigra Collin in<br />
association with <strong>the</strong> elm bark beetle Scolytus scolytus F. There is scope for much more<br />
research on o<strong>the</strong>r species <strong>of</strong> trees and <strong>the</strong>ir associated beetles and Medetera. Teskey<br />
( 1976) provides a useful summary <strong>of</strong> recent work on this <strong>to</strong>pic. Krivosheina ( 1974)<br />
describes and keys larvae <strong>of</strong> some Russian species including 5 that occur in Britain.<br />
The larvae <strong>of</strong> Thrypticus are plant-miners in <strong>the</strong> stems (fig. 1291) <strong>of</strong>Monocotyledons<br />
associated with wet environments, e.g. Eleocharis and Scirpus (Cyperaceae),<br />
Muhlenbergia and Phragmites (Gramineae) and ]uncus (Juncaceae). The female flies<br />
have a sharp sclerotised oviposi<strong>to</strong>r for piercing <strong>the</strong> food plant and <strong>the</strong> larvae and pupae<br />
have secondary structural changes associated with <strong>the</strong> non-carnivorous life-style in this<br />
specialised habitat (e.g. peripneustic larvae with reduced mouth parts and without anal<br />
lobes; pointed abdominal structures on pupae; no cocoon formed). No larvae <strong>of</strong> <strong>the</strong> 7<br />
<strong>British</strong> species have yet been described but Liibben (1908) has reared T. smaragdinus<br />
Gerstaecker and Dyte (1959) reviews work in America; ano<strong>the</strong>r compact research<br />
project awaiting investigation.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Cyrturella albose<strong>to</strong>sa Strobl are unknown but its only known<br />
<strong>British</strong> locality is Chippenham Fen, Cambs.<br />
Rhaphiinae. On <strong>the</strong> continent <strong>the</strong> larva <strong>of</strong> Rhaphium crassipes (Meigen) has been<br />
found in soil in a beech forest and Rh. elegantulum (Meigen) has been reared from<br />
a pupa found in sand at <strong>the</strong> edge <strong>of</strong> a pond (Lundbeck, 1912). None <strong>of</strong> <strong>the</strong> remaining<br />
23 species <strong>of</strong> Rhaphium occurring in Britain are known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> but<br />
McAlpine et al. ( 1981) illustrate a Nearctic species (fig. 217).<br />
Vaillant (1949) has described <strong>the</strong> larva <strong>of</strong> Syn<strong>to</strong>rmon zelleri Loew from wet rocks in<br />
company with Liancalus and Tachytrechus in France. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong><br />
remaining 11 <strong>British</strong> Syn<strong>to</strong>rmon species are unknown.<br />
Machaerium maritimae Haliday develops in <strong>the</strong> intertidal mud in estuaries (figs<br />
226--228).<br />
The larva <strong>of</strong> Systenus has a prominent pair <strong>of</strong> pro legs on <strong>the</strong> first abdominal segment<br />
(figs 219-220). Krivosheina (1973) describes and keys <strong>the</strong> larvae <strong>of</strong> S. scholtzii (Loew),<br />
S. pallipes Roser and S. tener Loew which are found in <strong>the</strong> accumulations <strong>of</strong> sap on tree<br />
trunks and in <strong>the</strong> humid rotten wood <strong>of</strong> tree holes. Lundbeck (1912) summarises <strong>the</strong><br />
o<strong>the</strong>r early rearing records which includeS. scholtzii from a beech fungus (by Verrall)<br />
and o<strong>the</strong>r <strong>British</strong> records.<br />
Achalcus melanotrichus Mik has been reared from rotten debris in lime, elm or horse<br />
chestnut trees. Bathycranium and Nema<strong>to</strong>proctus are unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Neurigoninae. The larva <strong>of</strong> Neurigona resembles Thrypticus in lacking anal lobes but<br />
unlike that genus is predaceous and has well developed mouthparts. Lundbeck ( 1912)<br />
gives a translation <strong>of</strong> Beling's brief description <strong>of</strong> <strong>the</strong> larva <strong>of</strong> N. quadrifasciata (F.)<br />
which was reared from soil under leaves in a forest <strong>of</strong> beech trees. The <strong>immature</strong> <strong>stages</strong><br />
<strong>of</strong> <strong>the</strong> 3 o<strong>the</strong>r <strong>British</strong> species are unknown.<br />
Diaphorinae. Dyte (1959) has reared Argyra argentella (Zetterstedt) and A . confinis<br />
(Zetterstedt) from mud, and <strong>the</strong> former from wet decaying sawdust near which dung<br />
had been dropped. Crisp & Lloyd (1954) reared A. leucocephala (Meigen) from mud<br />
but no larva has been adequately described. Dyte also notes that two American species<br />
<strong>of</strong> Diaphorus have been reared from human faeces but nothing <strong>of</strong> <strong>the</strong> life his<strong>to</strong>ries <strong>of</strong> <strong>the</strong><br />
4 <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus or <strong>of</strong> <strong>the</strong> genera Chrysotus (19 spp.) or Melanos<strong>to</strong>lus (I sp.)<br />
is known.<br />
69
Campsicneminae. Campsicnemus curvipes (Fallen) has been reared from mud (Crisp<br />
& Lloyd, 1954) and Nielsen et al. (1954) found larvae <strong>of</strong> what is probably C. armatus<br />
(Zetterstedt) (fig. 222) in <strong>the</strong> very wet soil <strong>of</strong> swamps and bogs near hot springs under<br />
mosses and grasses in Iceland. Nothing is known <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ries <strong>of</strong> <strong>the</strong> o<strong>the</strong>r 9<br />
<strong>British</strong> members <strong>of</strong> <strong>the</strong> genus or any <strong>of</strong> <strong>the</strong> remaining 9 genera <strong>of</strong> <strong>the</strong> subfamily.<br />
70
Key <strong>to</strong> families for final stage larvae <strong>of</strong> <strong>British</strong> Cyclorrhapha<br />
Larva distinctly sclerotized, flattened dorsoventrally, with prominent segmentation and<br />
with obvious long processes 2<br />
Larva less distinctly sclerotized or flattened dorsoventrally, with or without processes 4<br />
2 Tergal plates distinct (7 visible body segments), with thin striated lateral plates; first two<br />
segments and last segment with long filamen<strong>to</strong>us processes; inactive, in moist decaying<br />
vegetable material, leaf-litter, etc. (figs 229- 230) . Lonchopteridae (p. 76)<br />
Tergal plates distinct but without thin striated lateral margins and with lateral and dorsal<br />
processes not restricted <strong>to</strong> <strong>the</strong> first two and last segments . 3<br />
3 Posterior spiracles each on a short tuberculate dorsal process situated near <strong>the</strong> anterior<br />
margin <strong>of</strong> <strong>the</strong> last abdominal segment, this process terminating in three lobes each <strong>of</strong><br />
which bears a spiracular opening; sometimes active; in decaying organic material <strong>of</strong>ten <strong>of</strong><br />
animal origin (including carrion) and contaminated by urine or excrement (some Phoridae<br />
and Platypezidae may key out here but <strong>the</strong> lateral processes are usually unbranched and<br />
<strong>the</strong> posterior spiracles closer <strong>to</strong>ge<strong>the</strong>r) (figs 903- 908) . Fanniidae (p. 133)<br />
Posterior spiracles each ending on a short slender spiculate tuberculate process arising<br />
caudally on <strong>the</strong> last abdominal segment; in <strong>the</strong> sap <strong>of</strong> tree wounds (fig. 593) .<br />
Periscelididae (p. I 04)<br />
4 Posterior spiracles on a prominent large sclerotized plate; endoparasites <strong>of</strong> Hemiptera<br />
Homoptera; inactive (some Tephritidae may key out here but are not parasites and have<br />
fan-like anterior spiracles) (figs 247- 251) . Pipunculidae (p. 79)<br />
Posterior spiracles o<strong>the</strong>rwise . 5<br />
5 <strong>An</strong>terior spiracles more dorsally situated on <strong>the</strong> prothoracic segment and <strong>the</strong>refore<br />
closer <strong>to</strong>ge<strong>the</strong>r; mandible with longitudinal axis at oblique or right angles <strong>to</strong> rest <strong>of</strong><br />
cephalopharyngeal skele<strong>to</strong>n and usually with two or more pairs <strong>of</strong> equally sized teeth;<br />
mainly in leaf-mines (figs 650-668) .<br />
Agromyzidae (p. Ill)<br />
<strong>An</strong>terior spiracles more lateral in position, or absent; mandibles normally in same plane<br />
as rest <strong>of</strong> cephalopharyngeal skele<strong>to</strong>n and if <strong>to</strong>o<strong>the</strong>d <strong>the</strong> teeth are fewer and unequal in<br />
size 6<br />
6 Posterior spiracles each with numerous pore-like openings; endoparasites in insects or<br />
mammals 7<br />
Posterior spiracle each with three slit-like openings (occasionally with up <strong>to</strong> six similar<br />
openings) I 0<br />
7 Posterior spiracles usually dome-shaped, ei<strong>the</strong>r with circular wart-like protuberances each<br />
bearing several spiracular pores (some Tachinidae, e.g. Pelatachina, fig. 720, may key here,<br />
but are parasites <strong>of</strong> Lepidoptera), or with clusters <strong>of</strong> pores radiating from <strong>the</strong> ecdysal scar,<br />
or with a small stellar process near each spiracular plate, or with a pair <strong>of</strong> large laterally<br />
directed anal vesicles; endoparasites <strong>of</strong> bees and wasps; inactive (figs 282- 292) .<br />
Conopidae (p. 85)<br />
Posterior spiracles o<strong>the</strong>rwise . 8<br />
8 Openings <strong>of</strong> posterior spiracle arranged m three radiating groups; endoparasites <strong>of</strong><br />
Hemiptera-Heteroptera . . . Tachinidae (part) (p. 11 8)<br />
Openings <strong>of</strong> posterior spiracle arranged circularly but not extending completely round <strong>the</strong><br />
spiracular plate, <strong>the</strong> inner side <strong>of</strong> <strong>the</strong> plate more or less devoid <strong>of</strong> openings . 9<br />
9 Posterior spiracles on dorsal surface <strong>of</strong> a transverse cleft <strong>of</strong> <strong>the</strong> terminal abdominal segment<br />
and are occluded within <strong>the</strong> cavity when opposing surfaces are brought <strong>to</strong>ge<strong>the</strong>r;<br />
endoparasites <strong>of</strong> sheep and deer (figs 683-692) Oestridae (p. 115)<br />
Posterior spiracles on evenly rounded terminal extremity <strong>of</strong> body where <strong>the</strong>y are unprotected;<br />
endoparasites <strong>of</strong> cattle, horses and deer (figs 693- 702) Hypodermatidae (p. 116)<br />
I 0<br />
Posterior spiracles close <strong>to</strong>ge<strong>the</strong>r on a fused process <strong>of</strong> varying length, from short <strong>to</strong> an<br />
elongate retractile tube (figs 252-281) . Syrphidae (p. 80)<br />
Posterior spiracles not on a fused process and, if appearing fused, <strong>the</strong>n lacking spicules,<br />
tubercles or dense pubescence . 11<br />
11 Neck region <strong>of</strong> larva with an incomplete U-shaped collar or ring (figs 395, 648, r) . 12<br />
Neck region <strong>of</strong> larva without such a collar or ring . 13<br />
12 Larva more slender, anterior segments more obviously tapering <strong>to</strong>wards head; numerous<br />
and obvious papillae laterally and ventrally on body; free living, feeding on aphids,<br />
coccids, etc., active (figs 385- 395) Chamaemyiidae (p. 91)<br />
71
Larva short and broad, last segment broad and rounded, anterior segments not so tapering<br />
<strong>to</strong>wards head; papillae smaller and confined <strong>to</strong> anterior segments; in wax cells <strong>of</strong> hive bees<br />
feeding on pollen paste, inactive (figs 647-649) . Braulidae (p. 11 0)<br />
13 <strong>An</strong>terior spiracle simple, unbranched; larva oval in outline with branched lateral<br />
appendages or more or less parallel-sided, somewhat depressed, with or without short<br />
dorsal and lateral processes 14<br />
<strong>An</strong>terior spiracle branched or absent . 15<br />
14 Posterior spiracles with four openings arranged radially around <strong>the</strong> ecdysal scar (fig. 241 ); in<br />
fungi; inactive (figs 241- 246) Platypezidae (p. 78)<br />
Posterior spiracle with four openings arranged in two pairs, one behind <strong>the</strong> o<strong>the</strong>r (figs. 232);<br />
polyphagous saprophages; in fungi, dung, carrion, etc.; parasitic or predaceous on<br />
molluscs, spiders, millipedes, and o<strong>the</strong>r insects (figs 231- 240) Phoridae (p. 76)<br />
15 Margins <strong>of</strong> each posterior spiracular plate with a complete ring <strong>of</strong> branched setae; in<br />
decaying seaweed, etc. on beaches, active (figs 406-412) . Coelopidae (p. 93)<br />
Margins <strong>of</strong> posterior spiracular plates without setae or setae present as isolated groups or<br />
tufts . 16<br />
16 Posterior spiracles on short processes ending in a sharp apex (Diastatidae may key<br />
here) . 17<br />
Posterior spiracles o<strong>the</strong>rwise . 18<br />
17 <strong>An</strong>terior spiracles absent; aquatic, in plant stems, or as leaf-mines; or terrestrial and<br />
carnivorous; inactive or slightly active (figs 613-616)<br />
. Ephydridae (part) (Notiphilinae) (p. <strong>10</strong>6)<br />
<strong>An</strong>terior spiracles present; terrestrial, in leaf-mines; inactive (figs 633- 634) .<br />
Drosophilidae (part) (Scap<strong>to</strong>myza) (p. I 08)<br />
18 Posterior spiracles on <strong>the</strong> inner side <strong>of</strong> a short process; o<strong>the</strong>r processes or tubercles may<br />
occur on <strong>the</strong> last segment; in decaying organic material <strong>of</strong> animal origin; active (figs<br />
552- 563) . Piophilidae (part) (p. <strong>10</strong>1)<br />
Posterior spiracles not on inner side <strong>of</strong> a spiracular process . 19<br />
19 Posterior spiracles on elongated processes forming a pair <strong>of</strong> diverging branches from a<br />
cylindrical elongated base, or at least <strong>the</strong> branches are united at <strong>the</strong> base (in Canacidae,<br />
couplet 23, <strong>the</strong> spiracular branches are not forked or diverging but are on elongated<br />
processes); mouth-hooks with accessory sclerites (small sclerites below mouth-hooks) 20<br />
Posterior spiracles on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> last segment <strong>of</strong> <strong>the</strong> abdomen, or on short processes<br />
which are not united at <strong>the</strong> base; mouth-hooks with or without accessory sclerites<br />
(Limnophora, Muscidae, fig. 949, has <strong>the</strong> posterior spiracles on diverging processes which<br />
are united basally, mouth-hooks with accessory sclerites, but occurs among moss in<br />
running water) 27<br />
20 Mouth-hooks united by a sclerotized ventral arch (fig. 508, va); thoracic segments longer and<br />
narrower than <strong>the</strong> abdominal segments; dark coloured active larva with <strong>the</strong> body retractile,<br />
usually in ditches, ponds, etc., terrestrial, aquatic or semi-aquatic, feeding on snails<br />
and o<strong>the</strong>r molluscs (figs 475- 518) Sciomyzidae (p. 96)<br />
Mouth-hooks not united by a sclerotized arch; thoracic segments not longer and narrower<br />
than <strong>the</strong> abdominal segments; not dark-coloured, active, retractile larva . 21<br />
21 Spines or setae present on at least <strong>the</strong> posterior body segments, apart from any on <strong>the</strong><br />
segmental borders 22<br />
Cuticle smooth, without spines or setae on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> body segments, although<br />
tubercles may be present 25<br />
22 Posterior part <strong>of</strong> body obviously swollen, bulbous in shape; anterior spiracles with a more or<br />
less elongated cylindrical central axis from which arise lateral processes; in decaying<br />
organic material such as dung, carrion, etc.; active (figs. 447-474) . Sepsidae (p. 95)<br />
Posterior part <strong>of</strong> body not obviously swollen; anterior spiracles usually more fan-shaped,<br />
sometimes elongated . 23<br />
23 Each posterior spiracle at <strong>the</strong> end <strong>of</strong> an elongated cylindrical process which projects from a<br />
tubular siphon derived from <strong>the</strong> last abdominal segment; ventral pseudopods <strong>of</strong>ten<br />
present; aquatic on plants, <strong>of</strong>ten anchored by <strong>the</strong> posterior pseudopods, or in semi-aquatic<br />
habitats which are contaminated with organic material 24<br />
Each posterior spiracle much shorter, without a basal tubular siphon derived from <strong>the</strong> last<br />
segment; in decaying organic material, such as fallen leaves (in which <strong>the</strong>y may mine) or in<br />
birds' nests, occasionally phy<strong>to</strong>phagous in roots and stems (figs 396-405)<br />
Lauxaniidae (p. 92)<br />
72
24<br />
25<br />
26<br />
28<br />
29<br />
30<br />
31<br />
32<br />
33<br />
34<br />
35<br />
Posterior spiracular process not forked, spiracles united (figs. 645- 646) Canacidae (p. 1<strong>10</strong>)<br />
Posterior spiracular process forked; spiracles not united (figs. 618-624) .<br />
. Ephydridae (part) (p. <strong>10</strong>6)<br />
<strong>An</strong>terior spiracles with a basal cylindrical part from <strong>the</strong> distal end <strong>of</strong> which arise several finer<br />
filaments, or anterior spiracles absent; saprophagous, in decaying fruit, or fruit-based<br />
food products, or o<strong>the</strong>r vegetable material, dung, fungi, etc.; active (figs 635-638) .<br />
. Drosophilidae (part) (p. <strong>10</strong>8)<br />
<strong>An</strong>terior spiracles present but o<strong>the</strong>rwise formed 26<br />
<strong>An</strong>terior spiracles with a very long slender central axis from which arise short lateral<br />
processes; posterior spiracles on an elongated siphon or process; body narrow, with<br />
ventral pseudopods; in sap from tree wounds, etc.; inactive (figs 590--591) .<br />
Aulacigastridae (p. <strong>10</strong>4)<br />
<strong>An</strong>terior spiracles with a long but broader central process, <strong>the</strong> distal end broadened and<br />
bearing <strong>the</strong> spiracular openings; no lateral processes; body broader, with ventral<br />
pseudopods; in larval tunnels <strong>of</strong> beetles (Scolytidae, etc.); inactive (figs 579-585)<br />
Odiniidae (p. <strong>10</strong>3)<br />
Mouthparts usually with accessory sclerites (fig. 911 , ob, ar) by mouth-hooks; posterior<br />
spiracular slits usually straight or sharply bent about midpoint (figs 925, 930) (sinuous in<br />
Mesembrina, figs 913- 914); under bark, under moss, in soil, etc.; inactive<br />
Muscidae (part) (p. 134)<br />
Mouthparts without accessory sclerites by mouth-hooks (fig. 919); in doubtful cases <strong>the</strong>n<br />
posterior spiracles with strongly sinuous slits (figs 956, 960) . 28<br />
Posterior spiracles with three strongly sinuous slits whose axes lie approximately parallel <strong>to</strong><br />
<strong>the</strong> spiracular margins (figs 913, 916) Muscidae (Muscinae) (p. 134)<br />
Posterior spiracles with straight slits, or if openings are sinuous <strong>the</strong>n <strong>the</strong>ir axes lie tangential<br />
<strong>to</strong> <strong>the</strong> outer spiracular margin . 29<br />
Each posterior spiracle with an obvious spine or lobe . 30<br />
Each posterior spiracle without such a spine or lobe 32<br />
Cephalopharyngeal skele<strong>to</strong>n vestigial, unpigmented, except <strong>the</strong> small mandibles; in<br />
decaying wood <strong>of</strong> fallen trees; inactive (fig. 578) . Clusiidae (p. <strong>10</strong>3)<br />
Cephalopharyngeal skele<strong>to</strong>n normal and pigmented . 31<br />
Posterior spiracles with <strong>the</strong> slits short, oval in shape, and arranged almost at right angles <strong>to</strong><br />
each o<strong>the</strong>r:<br />
(a) in plant roots and stems (figs 361- 372) Psilidae (p. 90)<br />
(b) associated with trees Lonchaeidae (part) (p. <strong>10</strong>1)<br />
(c) in seaweed (figs 373- 379) . Helcomyzidae (part) (p. 91)<br />
Posterior spiracles with <strong>the</strong> openings radiating from <strong>the</strong> ecdysal scar at distinctly less than<br />
right angles, or irregularly or peripherally arranged:<br />
(a) in root-nodules <strong>of</strong> legumes or decaying organic material; inactive (figs 338- 352) .<br />
Micropezidae (p. 89)<br />
(b) in roots and bulbs (figs 336-337) Otitidae (part) (p. 88)<br />
(c) with bifid mouth-hooks in ]uncus, Carex, etc.; or mining leaves <strong>of</strong> water lilies (figs<br />
835- 838; 849- 851) . Scathophagidae (part) (Cordilura, Hydromyza) (p. 129)<br />
(d) in decaying wood (figs 607-609). . Diastatidae (p. <strong>10</strong>8)<br />
<strong>An</strong>terior spiracle bicornuate (fig. 359), posterior spiracles each on a distinct process (fig.<br />
356); saprophagous . . Tanypezidae (p. 90)<br />
<strong>An</strong>terior spiracle o<strong>the</strong>rwise, or if bicornuate (some Scathophagidae, fig. 840), <strong>the</strong>n posterior<br />
spiracles not on raised processes . 33<br />
Posterior spiracles united, <strong>the</strong> three spiracular openings <strong>of</strong> each weakly or strongly bent;<br />
endoparasitic in horses; inactive (figs 703- 711) . Gasterophilidae (p. 117)<br />
Posterior spiracles separate 34<br />
Posterior spiracles in a distinct deep depression (fig. 781); in decaying organic matter <strong>of</strong><br />
animal origin, carrion, etc.; active (figs 768, 785) . . Sarcophagidae (p. 125)<br />
Posterior spiracles not in a sunken depression 35<br />
Apex <strong>of</strong> each posterior spiracular process separated in<strong>to</strong> three, on each <strong>of</strong> which is an<br />
opening:<br />
(a) in excrement, carrion, or ants' nests (figs 639-644) .<br />
(b) in refuse in birds' nests (figs 586-589) .<br />
(c) plant feeders or preda<strong>to</strong>rs <strong>of</strong> root aphids (figs 669-682)<br />
Apex <strong>of</strong> each posterior spiracular process normal .<br />
. Milichiidae (p. 1<strong>10</strong>)<br />
Carnidae (p. <strong>10</strong>4)<br />
Chloropidae (part) (p. 113)<br />
36<br />
73
36 <strong>An</strong>terior spiracle with a more or less elongated central axis from which arise short or longer<br />
processes laterally; posterior spiracular openings short, elliptical; in carrion, dung, nests,<br />
etc., usually associated with decaying organic material (figs 519- 530) .<br />
. Sphaeroceridae (p. 99)<br />
Not as above . 37<br />
37 Posterior spiracular openings ra<strong>the</strong>r sinuous, all openings more or less parallel <strong>to</strong> each o<strong>the</strong>r<br />
and <strong>to</strong> <strong>the</strong> outer margin <strong>of</strong> <strong>the</strong> spiracle; mouth-hooks strongly curved near base, rest <strong>of</strong><br />
cephalopharyngeal skele<strong>to</strong>n united; under bark; inactive (figs 353- 354) .<br />
Megamerinidae (p. 90)<br />
Not as above . 38<br />
38 Posterior spiracles with three very short elliptical openings (some <strong>An</strong>thomyiidae may key<br />
here, but are included in o<strong>the</strong>r half <strong>of</strong> couplet) 39<br />
Posterior spiracles with more elongated, slit-like openings 40<br />
39 <strong>An</strong>terior spiracles fan-like:<br />
(a) phy<strong>to</strong>phagous, esp. in Gramineae (figs 669-682) Chloropidae (part) (p. 113)<br />
(b) predaceous on bark beetles or phy<strong>to</strong>phagous in plants (figs 531 - 538)<br />
Pallopteridae (p. I 00)<br />
(c) in water-loving Gramineae (figs 594-606) <strong>An</strong>thomyzidae (part) (p. <strong>10</strong>5)<br />
<strong>An</strong>terior spiracles with central axis elongated:<br />
(a) in Gramineae (figs 564-577) . Opomyzidae (p. <strong>10</strong>2)<br />
(b) in water-loving Gramineae (figs 594-606) <strong>An</strong>thomyzidae (part) (p. <strong>10</strong>5)<br />
40 Posterior spiracles on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> last segment, not at all raised on short processes 41<br />
Posterior spiracles on <strong>the</strong> end <strong>of</strong> more or less obvious, though always short, processes 42<br />
41 End <strong>of</strong> last abdominal segment with obvious tubercles arranged in a more or less circular<br />
sequence around <strong>the</strong> posterior surface; spiracular scar (but<strong>to</strong>n) usually in <strong>the</strong> ra<strong>the</strong>r<br />
narrow but strongly sclerotized spiracular border (peritreme); in decaying organic matter<br />
<strong>of</strong> animal origin, carrion, dung, wounds <strong>of</strong> animals, etc.; active maggots (figs 786-825) .<br />
Calliphoridae (p. 127)<br />
End <strong>of</strong>last segment with less obvious tubercles; spiracular border without <strong>the</strong> spiracular scar<br />
which is always situated inside <strong>the</strong> border; in plant stems, roots, fruits, flower-heads,<br />
sometimes causing mines or galls (figs 293- 314) Tephritidae (p. 86)<br />
42 Endoparasitic in insects, woodlice (lsopoda), etc 43<br />
Free living larvae 44<br />
43 Endoparasitic in insects (mainly) (figs 712- 755). Tachinidae (part) (p. 118)<br />
Endoparasitic in woodlice (figs 756-767) . Rhinophoridae (p. 124)<br />
44 Body, including last abdominal segment, without obvious papillae or tubercles:<br />
(a) in root nodules <strong>of</strong> legumes (Rivellia) ; in fungi , soil or dead wood (P/atys<strong>to</strong>ma) (figs<br />
315- 321) . Platys<strong>to</strong>matidae (p. 88)<br />
(b) in decaying vegetable matter or dung (figs 322- 337) . Otitidae (part) (p. 88)<br />
(c) under bark, in dung, plant roots and stems, conifer cones, fruits, etc. (figs 539- 547)<br />
. Lonchaeidae (part) (p. <strong>10</strong>1)<br />
(d) in fungi , carrion, dung, decomposing matter (figs 413- 446) .<br />
. Heleomyzidae (part) (p. 94)<br />
(e) in decaying seaweed on beaches (figs 373- 379) . Helcomyzidae (part) (p. 91)<br />
Body, including last segment, with a ring <strong>of</strong> papillae or tubercles around <strong>the</strong> spiracular field<br />
and with o<strong>the</strong>rs on o<strong>the</strong>r segments; cuticle may have spines or hairs 45<br />
45 <strong>An</strong>terior spiracles (fig. 638) consisting <strong>of</strong> long filaments situated at <strong>the</strong> tip <strong>of</strong> an <strong>of</strong>ten long<br />
basal stem; in decaying or fermenting organic matter (figs 625-638)<br />
Drosophilidae (part) (p. <strong>10</strong>8)<br />
<strong>An</strong>terior spiracles o<strong>the</strong>rwise . 46<br />
46 Slits in posterior spiracles small, at right angles <strong>to</strong> each o<strong>the</strong>r and well separated basally; in<br />
fungi, dung, carrion, rotting wood and o<strong>the</strong>r decomposing material (figs 413- 446).<br />
. Heleomyzidae (part) (p. 94)<br />
Slits in posterior spiracles larger, usually at less than 90° <strong>to</strong> each o<strong>the</strong>r and more<br />
approximated basally 47<br />
47 Cephalopharyngeal skele<strong>to</strong>n without a distinct sclerite (ventral arch) beneath <strong>the</strong> mouthhooks<br />
(figs 548-551 ); sucking blood <strong>of</strong> nestling birds .<br />
. Piophilidae (part) (Neottiophilinae) (p. <strong>10</strong>1)<br />
Cephalopharyngeal skele<strong>to</strong>n with a distinct (<strong>of</strong>ten sub-triangular) ventral sclerite 48<br />
74
48 <strong>An</strong>al fleshy lobes long, more equal in size, arranged with 8 evenly spaced around <strong>the</strong> discal<br />
plate, plus 2 more ventrally; in fungi, carrion (figs 380--384) . . Dryomyzidae (p. 91)<br />
<strong>An</strong>al lobes o<strong>the</strong>rwise . . . . . . . . . . . . . . . . . . . . . 49<br />
49 <strong>An</strong>al lobes usually separate and simple; in dung, plant stems or leaf mines in docks, reeds,<br />
water-lilies, orchids, etc. (figs 829- 854) . . . . . . Scathophagidae (part) (p. 129)<br />
<strong>An</strong>al lobes sometimes joined basally or individual lobes bifurcate; in stems or roots <strong>of</strong> plants,<br />
leaf-mines, fungi or rotting wood (figs 855- 878) . . . . . . <strong>An</strong>thomyiidae (p. 130)<br />
Families not included in <strong>the</strong> key<br />
Hippoboscidae (p. 139) and Nycteribiidae (p. 140) larvae (prepupae) pupate soon<br />
after <strong>the</strong> female deposits <strong>the</strong>m. Larvae have a large sclerotized spiracular plate and a<br />
distinct spiracular field and <strong>the</strong> spiracular openings are connected by branching tubular<br />
structures (fig. 967).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> following families are undescribed (see under each family<br />
for known habits):<br />
Acar<strong>to</strong>phthalmidae (in fungi, dung, carrion) (see comment below)<br />
Asteiidae (in fungi, reeds, wood-detritus)<br />
Camillidae ( in mammal nests, bird-guano)<br />
Chyromyidae (in bird and bat-guano)<br />
Stenomicridae (in water holding leaf-bases <strong>of</strong>Gramineae etc)<br />
Tethinidae (saline habitats, shores, dunes etc)<br />
Trixoscelidae (bird-guano)<br />
As this book goes <strong>to</strong> press a very important work on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
Cylorrhapha has appeared (Ferrar, 1987) and <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Acar<strong>to</strong>phthalmus<br />
bicolor Oldenberg have been described from carrion (Ozerov, 1987).<br />
75
Notes on families <strong>of</strong> Cyclorrhapha<br />
Series Aschiza<br />
Lonchopteridae<br />
(Figs: 229- 230, puparium 1149, pupa 1150)<br />
This small family has only 37 described world species, all belonging <strong>to</strong> <strong>the</strong> single<br />
genus Lonchoptera. Seven species occur in Britain <strong>of</strong> which two are known in <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong>. The larvae have been found under logs, among dead leaves and in<br />
decaying vegetable matter generally. They have also been found between <strong>the</strong> leaves <strong>of</strong><br />
Brussels sprouts and on <strong>the</strong> moist surface <strong>of</strong> swede turnips, especially during wet<br />
wea<strong>the</strong>r. The transport <strong>of</strong> vegetables coupled with par<strong>the</strong>nogenetic reproduction in <strong>the</strong><br />
adult may explain <strong>the</strong> cosmopolitan distribution <strong>of</strong> L.furcata (Fallen).<br />
The larva (figs 229- 230) is <strong>of</strong> a striking appearance, flattened ventrally with an<br />
arched back and long paired processes at each end. The larva <strong>of</strong> L. lutea Panzer has<br />
been described by Lubbock ( 1862) and de Meijere ( 1900) and that <strong>of</strong> L. furcata by<br />
Nielsen et al. (1954). No o<strong>the</strong>r species are known in <strong>the</strong> larval stage. Whit<strong>to</strong>n (1956)<br />
describes <strong>the</strong> respira<strong>to</strong>ry system, which is amphipneustic.<br />
Phoridae<br />
(Figs: larvae 231- 240; egg I 007; puparia 1151 ,1152, 1154; pupa 1153)<br />
The Phoridae (scuttle-flies) is a large family with about 3,000 species so far described<br />
throughout <strong>the</strong> world. The life-his<strong>to</strong>ries are poorly known but <strong>the</strong> larvae show a very<br />
wide range <strong>of</strong> habits from polyphagous saprophages and fungus feeders <strong>to</strong> specialised<br />
preda<strong>to</strong>rs, parasi<strong>to</strong>ids and true parasites. Some 300 species in 22 genera occur in<br />
Britain. The subfamilies are ill-defined even in <strong>the</strong> adult stage and no larval key is<br />
<strong>of</strong>fered but for convenience <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are discussed under <strong>the</strong> 3 subfamilies<br />
recognised in <strong>the</strong> Check List (Smith, 1976). Disney (1979, 1983) reviews <strong>the</strong> biology <strong>of</strong><br />
<strong>the</strong> <strong>British</strong> Phoridae and cites <strong>the</strong> original records for most <strong>of</strong> <strong>the</strong> comments below.<br />
Much careful work is needed in working out life-his<strong>to</strong>ries <strong>of</strong> this neglected but<br />
biologically important family now that good Handbooks <strong>to</strong> adults (Disney, 1983, 1989)<br />
are available.<br />
Aenigmatiinae. The single <strong>British</strong> genus Aenigmatias is known <strong>to</strong> parasitize <strong>the</strong><br />
pupae <strong>of</strong> ants, particularly wood ants (Formica spp.). The <strong>immature</strong> <strong>stages</strong> have not<br />
been adequately described and <strong>the</strong> identities <strong>of</strong> <strong>the</strong> three <strong>British</strong> species have been<br />
confused in recorded rearings.<br />
Me<strong>to</strong>piniinae. This subfamily includes <strong>the</strong> large genus Megaselia (figs 231- 233)<br />
with over 200 <strong>British</strong> species and a wide range <strong>of</strong> life-his<strong>to</strong>ries. Megaselia rufipes<br />
(Meigen) is <strong>the</strong> most frequently met <strong>of</strong> <strong>the</strong> polyphagous sa prophage species with M.<br />
giraudii (Egger) a close second (records <strong>of</strong> both species as parasi<strong>to</strong>ids are probably<br />
incorrect). The larvae <strong>of</strong> M. ha/terata (Wood) and M. nigra (Meigen) are known fungus<br />
feeders but o<strong>the</strong>r species occurring in fungus may be preda<strong>to</strong>rs or parasites <strong>of</strong> o<strong>the</strong>r<br />
fungivorous Diptera. M. aequalis (Wood) and M. ciliata (Zetterstedt) feed on slug eggs;<br />
M . melanocephala (von Roser), M. nasoni (Malloch) and M. pulicaria (Fallen) feed on<br />
spider eggs; M.fuscinervis (Wood) attacks snails ( Vitraea); M. brevicostalis (Wood) has<br />
been reared from deal snails (Helix and Cepaea). M. jlavicoxa (Zetterstedt) and<br />
M. obscuripennis Wood parasitize larval Sciaridae (Trichosia, Bradysia); M. paludosa<br />
(Wood) parasitizes Tipulidae larvae. The larvae <strong>of</strong> some non-<strong>British</strong> species <strong>of</strong><br />
Megaselia are parasitic on millipedes and predaceous on coccids.<br />
76
The two <strong>British</strong> Pseudacteon species are parasi<strong>to</strong>ids <strong>of</strong> adult worker ants; P.<br />
brevicauda Schmitz on Myrmica ruginodis Nylander and M . scabrinodis Nylander; P.<br />
f ormicarum on Lasiusflavus (F.), L. niger (L.), Formica sanguine a Latreille and possibly<br />
o<strong>the</strong>r genera.<br />
Phalacro<strong>to</strong>phora species parasitize <strong>the</strong> pupae <strong>of</strong> ladybird beetles (Coccinellidae).<br />
The <strong>immature</strong> <strong>stages</strong> are undescribed for <strong>the</strong> genera Beckerina, Gymnophora,<br />
Me<strong>to</strong>pina, Plectanocnema and Woodiphora. However, Me<strong>to</strong>pina occurs in buried<br />
carrion (see Smith, 1986b) and Woodiphora retroversa (Wood) has been associated with<br />
a Cossus (Lepidoptera) excavation in a tree (Disney, 1983). The genus Plas<strong>to</strong>phora <strong>of</strong><br />
<strong>the</strong> Check List is now included in Megaselia.<br />
Chonocephalus is a genus <strong>of</strong> about 40 species mainly found in <strong>the</strong> tropics and subtropics<br />
with 3 species in <strong>the</strong> Palaearctic Region. The larvae (fig. 234) <strong>of</strong> a few species<br />
have been reported in rotting fruits and rotting palm spadices and may thus come <strong>to</strong> <strong>the</strong><br />
notice <strong>of</strong> economic en<strong>to</strong>mologists. C. heymonsi S<strong>to</strong>bbe has occurred in a London<br />
hothouse.<br />
Phorinae. The larvae (fig. 235) <strong>of</strong> <strong>An</strong>evrina (4 <strong>British</strong> species) occur on vertebrate<br />
carrion and in <strong>the</strong> soil, particularly in mole nests.<br />
Borophaga incrassata (Meigen) (figs 236-237) parasitizes <strong>the</strong> larvae <strong>of</strong> Bibio marci L.<br />
(Bibionidae) (Morris, 1922) but <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> o<strong>the</strong>r 5 <strong>British</strong> species are<br />
unknown though B. irregularis (Wood) has been reared from pupae under ash logs.<br />
Chae<strong>to</strong>pleurophora larvae have been found in dead snails.<br />
Conicera tibialis Schmitz is <strong>the</strong> 'c<strong>of</strong>fin fly' that breeds in c<strong>of</strong>fined bodies and appears<br />
on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> ground above <strong>the</strong> body after a year or more. This and o<strong>the</strong>r species<br />
have also been reared from o<strong>the</strong>r carrion. In addition C. dauci (Meigen) has been reared<br />
from fungi. Fur<strong>the</strong>r information on <strong>the</strong> biology <strong>of</strong> Conicera in carrion, including<br />
human cadavers, is summarised by Smith (1986b).<br />
Some species <strong>of</strong> Diplonevra breed in carrion; D.funebris (Meigen) has been reared<br />
from wasps' ( Vespula) nests; D. pilosella Schmitz has been reared from a wounded<br />
earthworm; D. nitidula (Meigen) adults have been seen swarming over a dead<br />
earthworm, and have been reared from digested sewage sludge.<br />
The tropical and sub-tropical genus Dohrniphora contains over <strong>10</strong>0 species. One<br />
species, D. cornuta (Bigot), has been spread by man and is now well established in<br />
Britain. The larvae (fig. 239) are found in sewage, compost, dead insects and snails,<br />
rotting ships' cargoes <strong>of</strong> rice, bran, cow peas, etc., vertebrate carrion, etc. and will<br />
attack <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> o<strong>the</strong>r species that compete with it for food (e.g. Psychoda<br />
alternata in trickling filter sewage beds, Kloter et al., 1977).<br />
Gymnoptera longicostalis Schmitz breeds in bumble-bee nests (Bombus) and G.<br />
vitripennis (Meigen) in wasps' nests but both species have been reared from <strong>the</strong> same<br />
caterpiller <strong>of</strong> Cossus cossus (L.) (goat moth).<br />
The larva <strong>of</strong> Phora holosericea Schmitz has been found preying upon root aphids<br />
some 20-40 cm below <strong>the</strong> soil surface (in Russia, Yarkulov, 1972). Apparently each<br />
larva consumes 80-85 aphids during its development. Nothing is known <strong>of</strong> <strong>the</strong> lifehis<strong>to</strong>ries<br />
<strong>of</strong> <strong>the</strong> remaining 8 species <strong>of</strong> Phora. Early records <strong>of</strong> P. aterrima F. from<br />
carrion are misidentifications (<strong>of</strong> adults) <strong>of</strong> Conicera.<br />
Larvae <strong>of</strong> Spiniphora live in dead snails but S. bergenstammi (Mik) (fig. 240) is<br />
frequently found by environmental health <strong>of</strong>ficers in milk bottles. The puparia (fig.<br />
1154) are so firmly cemented <strong>to</strong> <strong>the</strong> inside <strong>of</strong> <strong>the</strong> bottle that <strong>the</strong>y can survive <strong>the</strong> normal<br />
washing process. S. bergenstammi has also been reared from an old nest <strong>of</strong> a blackbird.<br />
Several species <strong>of</strong> Triphleba breed in vertebrate carrion; T. antricola (Schmitz) also<br />
breeds in bat dung in caves; T. lugubris (Meigen) breeds in wasps' nests; T. minuta (F.)<br />
breeds in <strong>the</strong> fungus Gymnopilus junonius (Fries) ( = Pholiota spectabilis Gillet); T.<br />
gracilis (Wiedemann) has been reared from puparia found under <strong>the</strong> bark <strong>of</strong> rotting<br />
larch and spruce logs.<br />
77
The life-his<strong>to</strong>ry <strong>of</strong> Hypocera is unknown. The genus Citrago is now included in<br />
Triph/eba.<br />
Platypezidae<br />
(Figs: larvae 241- 246, cocoon 1156, puparium 1157)<br />
Some 200 species <strong>of</strong> Platypezidae (flat-footed flies) have been described throughout<br />
<strong>the</strong> world. In Britain <strong>the</strong>re are 31 species (including Ate/estus, see below) in 12 genera<br />
representing three subfamilies. All those Platypezidae known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong><br />
are fungus feeders. Kessel et al. ( 1973) survey what is known <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and<br />
key <strong>10</strong> genera.<br />
Opetiinae. The <strong>immature</strong> <strong>stages</strong> and life-his<strong>to</strong>ries <strong>of</strong> this subfamily are unknown.<br />
Opetia nigra Miegen occurs in woodlands though <strong>the</strong> female is rarely seen. The genus<br />
probably belongs in <strong>the</strong> Empidoidea and discovery <strong>of</strong> <strong>the</strong> larva may resolve this problem<br />
(Disney, 1987). The three tiny species <strong>of</strong> Microsania are rarely seen except in <strong>the</strong><br />
smoke <strong>of</strong> bonfires <strong>to</strong> which <strong>the</strong>y are particularly attracted. The two species <strong>of</strong> Ate/estus<br />
are usually beaten from <strong>the</strong> foliage <strong>of</strong> trees in <strong>the</strong> spring but this genus has recently been<br />
placed in a separate family, <strong>the</strong> Atelestidae (Chvala, 1983). Some rearing data are<br />
provided by Chandler ( 1973, and in Stubbs & Chandler, 1978) and Brindle ( 196le) keys<br />
<strong>the</strong> few <strong>British</strong> species known in <strong>the</strong> larval <strong>stages</strong>.<br />
Callomyiinae. The striking larvae <strong>of</strong> Callomy ia amoena Meigen (fig. 245) feed on <strong>the</strong><br />
surface <strong>of</strong> bark encrusting fungi (Corticium spp.) in damp situations. The host fungi <strong>of</strong><br />
<strong>the</strong> o<strong>the</strong>r 3 species in <strong>the</strong> genus are not known. Members <strong>of</strong> <strong>the</strong> genus Agathomyia (fig.<br />
246) appear <strong>to</strong> develop internally in <strong>the</strong> <strong>to</strong>ugher Polyporaceae. A . unico/or Oldenberg<br />
and A.fal/eni (Zetterstedt) have both been reared from Bjerkandera ( Poly porus) adusta<br />
Fries and A. antennata (Zetterstedt) probably develops in Coria/us versicolor Fries,<br />
which <strong>the</strong> adults frequent. The host fungi <strong>of</strong> <strong>the</strong> remaining 6 <strong>British</strong> species <strong>of</strong><br />
Agathomyia are unknown.<br />
Platypezinae. Pro<strong>to</strong>clythia species attack s<strong>of</strong>t fungi, chiefly <strong>the</strong> honey fungus<br />
Armil/aria me/lea Quelet from which both <strong>British</strong> species (P. modest a (Zetterstedt) and<br />
P. rufa (Meigen)) have been reared. Two <strong>of</strong> <strong>the</strong> four <strong>British</strong> species <strong>of</strong> Platypeza (figs<br />
241- 244) also breed in <strong>the</strong> honey fungus but in addition P.fasciata Meigen has been<br />
reared from a puff-ball (Lycoperdon py riforme Schaeffer) and a Boletus sp. Dusek<br />
(1962) gives a good illustrated description <strong>of</strong> <strong>the</strong> larva and puparium <strong>of</strong> P.fasciata.<br />
Plesioclythia dorsalis (Meigen) breeds in field mushrooms (Agaricus spp.) but on <strong>the</strong><br />
continent it has also been reared from <strong>the</strong> giant puff-ball (Langermannia gigantea<br />
(Persoon) Rostkovius) and <strong>the</strong> cep (Boletus edulis Bulliard). I have also seen <strong>the</strong>m<br />
in boxes <strong>of</strong> chocolates perhaps attracted by a fungal 'bloom' or more likely having<br />
wandered from nearby mushrooms in searching for a pupation site!<br />
Paraplatypeza atra (Meigen) has been reared from Pluteus cervinus Quelet.<br />
Orthovenafurcata (Fallen) breeds in Polyporus squamosus Fries, and adults have also<br />
been seen on or near o<strong>the</strong>r fungi (Pleurotus sapidus Sacc. and C/i<strong>to</strong>pilus prunulus<br />
Quelet).<br />
Polyporivora infumata (Haliday) and P. picta Meigen have been reared from Coriolus<br />
versicolor.<br />
In spite <strong>of</strong> <strong>the</strong>se rearings very few larval descriptions or studies <strong>of</strong> host fungus<br />
specificity have been published and <strong>the</strong>re is scope for a research project here, perhaps<br />
coupled with o<strong>the</strong>r fungivorous Diptera, e.g. Myce<strong>to</strong>philidae. Peter Chandler will be<br />
describing <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> most <strong>of</strong> <strong>the</strong> above in his forthcoming volume in <strong>the</strong><br />
Fauna ent. Scand. series.<br />
78
Pipunculidae<br />
(Figs: larvae 247- 25 I, puparium I I 55)<br />
The family Pipunculidae contains about 600 species throughout <strong>the</strong> world. In Britain<br />
74 species in 8 genera have so far been recorded but <strong>the</strong> family needs fur<strong>the</strong>r careful<br />
study.<br />
The larvae <strong>of</strong> pipunculids are endoparasites <strong>of</strong> Hemiptera-Homoptera, especially<br />
Cicadellidae (leaf-hoppers), Cercopidae (frog-hoppers) and Delphacidae. The biology<br />
<strong>of</strong> <strong>the</strong> family is poorly known and very few larval descriptions have been published, but<br />
classic work by Perkins (1905) in Hawaii has established <strong>the</strong>ir value in <strong>the</strong> biological<br />
control <strong>of</strong> <strong>the</strong> sugarcane leaf-hopper (Perkinsiella saccharicida Kirkaldy). O<strong>the</strong>r<br />
economically important leaf-hoppers are attacked by Pipunculidae in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong><br />
world. Except for Eudorylas each genus is confined <strong>to</strong> a single host-family and many<br />
species are host specific, though some may range over several genera. Clausen (1940),<br />
Coe (1966) and Askew (1971) review <strong>the</strong> biology <strong>of</strong> <strong>the</strong> family. Much careful rearing,<br />
coupled with a refined taxonomic study <strong>of</strong> <strong>the</strong> adults, is badly needed in this family<br />
which presents problems similar <strong>to</strong> those in <strong>the</strong> parasitic Hymenoptera.<br />
Three subfamilies are represented in Britain.<br />
Chalarinae. The genus Chalarus confines its attentions <strong>to</strong> Cicadellidae <strong>of</strong> <strong>the</strong><br />
subfamily Typhlocybinae. Several <strong>of</strong> <strong>the</strong> 8 <strong>British</strong> species have been reared but early<br />
records are undoubtedly marred by misidentifications. Wal<strong>of</strong>f (I 97 5) records C. spur ius<br />
Falh!n from Eupteryx notata Curtis.<br />
Verrallia species (fig. 247) confine <strong>the</strong>ir attentions <strong>to</strong> <strong>the</strong> family Cercopidae. Unlike<br />
o<strong>the</strong>r Pipunculidae this genus appears <strong>to</strong> attack adult Homoptera <strong>to</strong> <strong>the</strong> exclusion <strong>of</strong><br />
<strong>the</strong> nymphal <strong>stages</strong>. V. aucta Fallen has been reared from <strong>the</strong> common <strong>British</strong> froghoppers<br />
Philaenus spumarius L. and Neophilaenus lineatus (L.). Whittaker (1969) and<br />
Wal<strong>of</strong>f(I 975) have studied <strong>the</strong> biology <strong>of</strong> V. aucta in <strong>the</strong>se two frog-hopper species and<br />
what is probably V. se<strong>to</strong>sa Verrall in Neophilaenus campestris (Fallen). V. pilosa<br />
Zetterstedt has been reared from a puparium found in a bird's nest, probably from a<br />
parasitized frog-hopper that had strayed or fallen in. No reliable rearing data are<br />
available for <strong>the</strong> remaining 3 <strong>British</strong> species.<br />
Nephrocerinae. Nephrocerus flavicornis Zetterstedt is <strong>the</strong> largest <strong>British</strong> pipunculid<br />
which has led <strong>to</strong> some speculation on <strong>the</strong> identity <strong>of</strong> <strong>the</strong> homopteran host capable <strong>of</strong><br />
sustaining its larva. Early in <strong>the</strong> century, when <strong>the</strong> only three <strong>British</strong> records <strong>of</strong> <strong>the</strong><br />
species were from <strong>the</strong> New Forest it was thought that <strong>the</strong> rare and restricted New<br />
Forest cicada (Cicadetta montana (Scopoli)) was <strong>the</strong> probable host. Subsequent<br />
records <strong>of</strong> N.flavicornis suggest that our second largest homopteran, Ledra aurita (L.),<br />
may be <strong>the</strong> true host since it shares a predilection for oak woods and is fairly widely<br />
distributed from <strong>the</strong> New Forest <strong>to</strong> Suffolk.<br />
Pipunculinae. Tomosvaryella ( = Alloneura) has seven <strong>British</strong> species and Wal<strong>of</strong>f<br />
(I 975) has reared three <strong>of</strong> <strong>the</strong>m from Cicadellidae: T. sylvatica (Meigen) from Jassargus<br />
pseudocellaris (Flor), Errastunus ocellaris (Fallen) and Arthaldeus pascuellus (Fallen);<br />
T. palliditarsis (Collin) from Diplocolenus abdomina/is (F.); and T. kuthyi (Aczel) from<br />
Psammotettix confinis (Dahlbom). On <strong>the</strong> continent Coe (1966) records T. sylvatica<br />
from Arthaldeus pascuellus and an unidentified Tomosvaryella sp. from Opsius sp.<br />
(both hosts are Cicadellidae).<br />
The only rearing records <strong>of</strong> Dorylomorpha (I I <strong>British</strong> species) are D. rujipes Meigen<br />
from Cicadula quadrinotata (F.) and D. xanthopus (Thomson) from an adult<br />
Psammotettix conjinis (Wal<strong>of</strong>f, I 975).<br />
The genus Pipunculus contains seven species including two <strong>of</strong> <strong>the</strong> commonest<br />
<strong>British</strong> members <strong>of</strong> <strong>the</strong> family, P. campestris Latreille and P. thomsoni Becker. Never<strong>the</strong>less<br />
breeding records are few. In Germany P. campestris has been reared from<br />
79
<strong>the</strong> Cicadellidae Macrosteles laevis Ribaut, Arthaldeus pascuel/us and Cicadula<br />
quadrinotata (Coe, 1966). Wal<strong>of</strong>f (1975) adds Eusce/is p/ebejus (Fallen), Psammotettix<br />
confinis and Elymana su/phurel/a (Zetterstedt) as hosts <strong>of</strong> P. campestris and also records<br />
P. thomsoni from Elymana su/phurel/a.<br />
Cephalops species (13 <strong>British</strong>) select Delphacidae as hosts and rearing records are<br />
comparatively numerous. Coe ( 1966) records C. semifumosus (Kowarz) from Ditropis<br />
pteridis Boheman and Conomelus anceps Germar; C. oberon Coe from D. pteridis;<br />
C. subultimus Collin from D. pteridis; and (from Germany) C. ultimus (Becker) from<br />
Eurysa lineata Perris. Wal<strong>of</strong>f (1975) records C. curtifrons Coe from Stenocranus<br />
minutus (F.) and C. semifumosus from <strong>the</strong> two hosts recorded by Coe (1966) and<br />
in addition Javesel/a pellucida (F.), Megame/odes venosus (Germar), Laodelphax<br />
elegantulus (Boheman), Dicranotropis hamata (Boheman), Javesel/a sp., and<br />
De/phacodes fairmarei (Perris). Rothschild (1964) describes <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong><br />
Cephalops semifumosus Kowarz (fig. 248) and May ( 1979) those <strong>of</strong> C. curtifrons Coe.<br />
The latter appears <strong>to</strong> be specific <strong>to</strong> Stenocranus minutus in which 35% parasitism may<br />
occur.<br />
Eudory/as species (21 <strong>British</strong>) (fig. 249) parasitize Cicadellidae. Wal<strong>of</strong>f (1975)<br />
discusses <strong>the</strong> biology and lists hosts (mostly Psammotettix confinis, Errastunus ocellar is<br />
and Arthaldeus pascuel/us) for E.fascipes (Zetterstedt), E.jenkinsoni Coe (=E. obliquus<br />
Coe), E. obscurus Coe, E. subfascipes Collin and E. subterminalis Collin. Coe (1966)<br />
lists some rearing records <strong>of</strong> Eudory/as from named cicadellid hosts in Germany.<br />
Syrphidae<br />
(Figs: larvae 252- 281, eggs I 008- <strong>10</strong>12, puparia 1158- 1164,<br />
pupa 1162, leaf mines 1292)<br />
About 5,000 species <strong>of</strong> Syrphidae (hover-flies) have been described throughout <strong>the</strong><br />
world. Of <strong>the</strong>se some 242 species in 68 genera occur in Britain.<br />
The larvae <strong>of</strong> some Syrphidae, especially aphidophagous species <strong>of</strong> <strong>the</strong> subfamily<br />
Syrphinae and <strong>the</strong> so called rat-tailed maggots (Eristalini), are quite well known but<br />
elsewhere knowledge is very irregular.<br />
The following key <strong>to</strong> subfamilies, and tribes <strong>of</strong>Milesiinae, is based on Hartley (1961)<br />
with indications where <strong>the</strong> extra tribes recognised in <strong>the</strong> Check List (Smith, 1976) are<br />
<strong>the</strong>re included. Notes on genera and species follow <strong>the</strong> tribal arrangement in <strong>the</strong> Check<br />
List. Keys <strong>to</strong> some genera and species for larvae with descriptions are provided by<br />
Dixon (1960) and Hartley (1961). For substantial, well illustrated, work on aphidophagous<br />
species see also Bhatia (1939) and Scott (1939). The continental works <strong>of</strong>Dusek &<br />
Laska (1967) and Goeldlin de Tiefenau (1974) are also very useful. Wood inhabiting<br />
species are keyed (in Russian) by Krivosheina & Mamaev (1967). O<strong>the</strong>r work is cited<br />
below. Stubbs & Falk (1983) provide a useful summary <strong>of</strong> habitats <strong>of</strong> <strong>the</strong> early <strong>stages</strong><br />
and key <strong>the</strong> adults.<br />
<strong>Part</strong>ial key <strong>to</strong> subfamilies and tribes <strong>of</strong> larval Syrphidae<br />
(Modified from Hartley, 1961)<br />
With piercing mouthparts. Posterior spiracles sessile, slits more or less radial and usually<br />
straight on slight ridges (except Xanthogramma where slits are convoluted but still<br />
distinctly radial); prothoracic spiracles very small. Prolegs vestigial or absent, without<br />
crochets; body <strong>of</strong>ten flattened and usually pointed anteriorly; usually with colour pattern<br />
which is lost on death. Predaceous, mainly on aphids (includes Syrphini, Chryso<strong>to</strong>xini,<br />
Bacchini, Melanos<strong>to</strong>matini and Paragini. Pipizini also key out here) . . Syrphinae<br />
Without piercing mouthparts. Posterior spiracles on a telescopic breathing tube; or if sessile<br />
with <strong>the</strong> slits basically more or less circumferential, usually convoluted and sometimes<br />
subdivided or as numerous small apertures, not on raised ridges; prothoracic spiracles<br />
80
present or absent. Pro legs with crochets present or absent. Shape varied; never with colour<br />
pattern but sometimes with brownish cuticle . . . Milesiinae 2<br />
2 <strong>An</strong>terior end pointed. Posterior spiracles with slits subdivided and radial. Prolegs well<br />
developed with few s<strong>to</strong>ut crochets. In nests <strong>of</strong> bees and wasps (figs 271- 276) <strong>Vol</strong>ucellini<br />
<strong>An</strong>terior end truncate or rounded. Posterior spiracles with slits undivided or if divided <strong>the</strong>n<br />
<strong>the</strong> pro legs are absent . . . . . . . . . . . . . . 3<br />
3 Body more or less hemispherical and flattened ventrally; pro- and mesothorax very reduced<br />
and only visible ventrally. Prolegs absent; pubescence usually as a lateral fringe; dorsal<br />
surface reticulated; metapneustic. In ants' nests (figs 269- 270) . . . Microdontini<br />
Body more or less cylindrical, pro- and mesothorax well developed and similar in size <strong>to</strong><br />
o<strong>the</strong>r segments . . . . . . . . . . . . . . . . . . . . 4<br />
4 Abdomen abruptly tapering <strong>to</strong> a long telescopic breathing tube; anterior spiracles elongate<br />
and retractile.ln stagnant foul water, liquid manure, rock pools, etc. (fig. 281) Eristalini<br />
Abdomen gradually tapering or truncate; if forming a distinct tail <strong>the</strong>n <strong>the</strong> anterior spiracles<br />
are squat and not retractile . . . . . . . . . . . . 5<br />
5 Metapneustic. Small white larvae (less than <strong>10</strong> mm) with long pubescence posteriorly;<br />
posterior spiracles convex or stylet-like on telescopic breathing tube. Larvae aquatic,<br />
piercing plant roots <strong>to</strong> obtain oxygen (figs 258- 259) ( Chrysogaster) . . . . . .<br />
. . . . . . . . . . . . . . . . . . . . . . Chrysogasterini (part)<br />
Amphipneustic (but anterior spiracle <strong>of</strong>ten very difficult <strong>to</strong> see). Larva o<strong>the</strong>rwise . . 6<br />
6 Body short (less than 8 mm) and flattened with abdomen gradually tapering posteriorly but<br />
not forming a distinct tail; prolegs and crochets present. Under wet decaying bark<br />
(Sphegina). In wet decaying manure etc (Neoascia) .<br />
Chrysogasterini (part)<br />
Body longer (more than <strong>10</strong> mm) and more or less cylindrical or if short and flattened <strong>the</strong>n<br />
truncate posteriorly and without prolegs . . . . . . . . . . . . . 7<br />
7 Abdominal pro legs large and partially fused in segmental pairs, with crochets <strong>of</strong> each pair<br />
confluent . . . . . . . . . . . . . . . . . . . . . Callicerini<br />
Prolegs separate or absent, crochets never confluent (includes Cheilosiini, Eumerini,<br />
Merodontini, Sericomyiini, Xylotini, Myolepta, Hammerschmidtia, Cheilosia) . . . .<br />
. . . . . . . . . . . . . . . . . . . Milesiini (s.lat.)<br />
Syrphinae<br />
Syrphini. The majority <strong>of</strong> genera and species in this tribe feed on aphids. The<br />
common Syrphus ribesii (L.) (fig. 253), S. vitripennis Meigen and S. <strong>to</strong>rvusOsten Sacken<br />
are widely polyphagous, eating a great variety <strong>of</strong> aphids and occur on many trees,<br />
shrubs and herbs. O<strong>the</strong>r genera with similarly polyphagous species include Epistrophe<br />
(fig. 254) (see Ro<strong>the</strong>ray, 1986b) Dasysyrphus (fig. 255), Episyrphus, Leucozona,<br />
Metasyrphus (s.s.), Scaeva and Sphaerophoria. Episyrphus balteatus (De Geer) is found<br />
abundantly on cabbages and would be a good species for use in demonstration and<br />
teaching. Some are more specialist feeders, e.g. Metasyrphus s.g. Lapposyrphus on<br />
conifer aphids (Adelges); Melangyna cincta Fallen on Phyllaphisfagi on Fagus silvatica.<br />
The rare Eriozona syrphoides (Fallen) has been found <strong>to</strong> feed on <strong>the</strong> aphid Cinaria<br />
pineae (Panzer) in established (40 years old) spruce monocultures in Czechoslovakia<br />
(Kula, 1983). It would be a useful study <strong>to</strong> rear Syrphini and identify <strong>the</strong>ir aphid prey <strong>to</strong><br />
establish <strong>the</strong> degree <strong>of</strong>specificity in habitat and in feeding, as Dusek & Laska (1966)<br />
have done in Czechoslovakia. The eggs, host range and oviposition behaviour have<br />
been studied by A. E. F. Chandler ( 1968). Some, e.g. Sphaerophoria, feed on o<strong>the</strong>r small<br />
Homoptera as well as Aphididae. Parasyrphus nigritarsis (Zetterstedt) attacks <strong>the</strong> eggs<br />
and larvae <strong>of</strong> certain chrysomelid beetles (Schneider, 1953); o<strong>the</strong>r members <strong>of</strong> <strong>the</strong><br />
genus are unknown as larvae and would well repay investigation. Papers by Ro<strong>the</strong>ray<br />
( 1986- still appearing) should be studied.<br />
The larval habits <strong>of</strong> some genera are unknown, but are probably aphidophagous, e.g.<br />
X anthogramma has been found under turf and s<strong>to</strong>nes and in ants' nests (Lasius spp.).<br />
Chryso<strong>to</strong>xini. Chryso<strong>to</strong>xum larvae appear <strong>to</strong> be <strong>of</strong> <strong>the</strong> aphidophagous type but <strong>the</strong><br />
feeding habits are unknown. They have been found under compost heaps, s<strong>to</strong>nes, etc.<br />
81
and Dixon (1960) reared C. verralli Collin from a larva found in an ants' nest (Lasius<br />
niger (L.)).<br />
Bacchini. Larvae <strong>of</strong> Baccha have been found feeding among colonies <strong>of</strong> aphids:<br />
Uromelanjaceae (L.) on Centaurea scabiosa L. and Brachycaudina napelli (Schrank) on<br />
Aconitum.<br />
Melanos<strong>to</strong>matini. Lucchese (1942) describes <strong>the</strong> larva <strong>of</strong> Xanthandrus comtus<br />
(Harris) which attacks caterpillars <strong>of</strong> Acroclita naevana Huebner and larvae <strong>of</strong> o<strong>the</strong>r<br />
small 'quasi-gregarious' <strong>to</strong>rtricid moths.<br />
Melanos<strong>to</strong>ma species normally feed on aphids but M. mellinum (L.) has also been<br />
recorded attacking <strong>to</strong>rpid adults <strong>of</strong> <strong>the</strong> flies Musca domestica L. and Paregle cinerella<br />
(Fallen) on umbels <strong>of</strong> Daucus carota L. When deprived <strong>the</strong> same species has been<br />
reported <strong>to</strong> take '<strong>to</strong>rtrix' larvae as prey. In <strong>the</strong> U.S.A. a non-<strong>British</strong> species has<br />
been reared on both aphids and decomposing chickweed, <strong>the</strong> latter being <strong>the</strong> more<br />
successful!<br />
Platycheirus species are aphid feeders. Dixon (1960) describes <strong>the</strong> larvae <strong>of</strong> several<br />
species, but gives no information on plant hosts or aphid food. P. scutatus (Meigen)<br />
is common on pota<strong>to</strong>es, cabbages, etc. and P. peltatus (Meigen) shows a strong<br />
preference for brassicas. P. ( Pachysphyria) ambiguus Fallen appears <strong>to</strong> specialize on<br />
aphids which curl up <strong>the</strong> leaves <strong>of</strong> Prunus (Dusek & Laska, 1966).<br />
Lundbeck (1916) described <strong>the</strong> larva and puparium <strong>of</strong> Pyrophaena granditarsa<br />
Forster and <strong>the</strong> puparium <strong>of</strong> P. rosarum (F.), both <strong>of</strong> which he found in flood refuse in<br />
fens (in Denmark). The feeding habits remain unknown but <strong>the</strong>y are thought <strong>to</strong> be<br />
aphidophagous.<br />
Paragini. The larvae <strong>of</strong> Paragus species (fig. 256) are aphidophagous.<br />
Milesiinae<br />
Pipizini. The larvae <strong>of</strong> Pipiza species (fig. 257) seem <strong>to</strong> prefer aphids which secrete a<br />
waxy flocculence, e.g. woolly aphid (Eriosoma). Pipizella larvae confine <strong>the</strong>ir attention<br />
<strong>to</strong> subterranean aphids feeding on <strong>the</strong> roots <strong>of</strong> plants.<br />
Heringia heringi (Zetterstedt) attacks <strong>the</strong> blister gall-forming aphid Schizoneura<br />
lanuginosa Hartig on elm (especially Ulmus procera Salisbury) and has been reared<br />
from an aphid gall on Salix, but <strong>the</strong> <strong>immature</strong> <strong>stages</strong> have not been adequately<br />
described.<br />
The larvae <strong>of</strong> Neocnemodon usually feed on aphids <strong>of</strong> <strong>the</strong> family Pemphigidae (e.g.<br />
Eriosoma) on deciduous trees, but N. latitarsis (Egger) and N. vitripennis (Meigen) have<br />
been found attacking <strong>the</strong> woolly aphid Dreyfusia piceae (Ratzeburg) (Adelgidae) on<br />
silver fir (Abies alba Miller).<br />
Triglyphus primus Loew larvae have been found feeding on aphids (Cryp<strong>to</strong>siphum<br />
artemisiae Buck<strong>to</strong>n, =gallarum Kaltenbach) on Artemisia vulgaris L.<br />
Cheilosiini. Cheilosia larvae are phy<strong>to</strong>phagous. Smith (1979b) summarises <strong>the</strong> plant<br />
associations which are with <strong>the</strong> stem bases and roots <strong>of</strong> higher plants (especially<br />
Compositae) or leaf mines in Crassulaceae and some fungi. Some additional information<br />
is provided by Stubbs (1983) and Ro<strong>the</strong>ray (1988a,b and in prep.) but<br />
host-plant information is still lacking for 20 <strong>of</strong> <strong>the</strong> 32 species occurring in Britain.<br />
The leaf-mining species is C. semifasciata Becker (fig. 1292) which attacks Umbilicus<br />
rupestris (Salisbury) Dandy (wall pennywort) and Sedum telephium L. (orpine). The<br />
species in fungi are closely related, e.g. C. scutellata (Fallen) and C. soror Zetterstedt<br />
(from truffles); C. scutellata and C.longula (Zetterstedt) in Boletus and allies (Leccinum,<br />
Suillus and Gyroporus).<br />
82
The larvae <strong>of</strong> Portevinia maculata (Fallen) live in <strong>the</strong> underground portion <strong>of</strong> <strong>the</strong> leaf<br />
bases <strong>of</strong> Allium ursinum L. (ramsons or wild garlic) (Dr M. C. D . Speight, pers. comm.).<br />
The common Rhingia campestris Meigen breeds in cow dung but nothing is known <strong>of</strong><br />
<strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> rare Rh. rostrata (L.).<br />
The larvae <strong>of</strong> Ferdinandea cuprea (Scopoli) have been found in <strong>the</strong> sap <strong>of</strong> wounds on<br />
tree trunks especially trees infested with <strong>the</strong> goat moth (Cossus cossus L.) around <strong>the</strong><br />
roots <strong>of</strong> which puparia have been found. It seems likely that <strong>the</strong> much rarer F. ruficornis<br />
(F.) breeds in similar situations.<br />
The early <strong>stages</strong> <strong>of</strong> Chamaesyrphus are unknown but should be sought in <strong>the</strong><br />
Caledonian pine woods where <strong>the</strong> adults occur among Calluna.<br />
Chrysogasterinae. The larvae <strong>of</strong> Myolepta resemble <strong>the</strong> 'rat-tailed' maggots <strong>of</strong> <strong>the</strong><br />
Eristalini but with a shorter anal siphon or 'tail'. They occur in wet tree rot-holes <strong>of</strong><br />
beech and poplar.<br />
The larvae <strong>of</strong> Chrysogaster are aquatic or semi-aquatic. Some species, e.g. C. hirtella<br />
Loew (figs 258- 259), have <strong>the</strong> posterior spiracles adapted as a stylet for piercing aquatic<br />
plants in order <strong>to</strong> obtain oxygen but o<strong>the</strong>r species do not, e.g. C. solstitialis (Fallen)<br />
(fig. 260).<br />
Orthonevra larvae (fig. 261) are found in mud containing decaying wood and o<strong>the</strong>r<br />
vegetation at <strong>the</strong> edge <strong>of</strong> ponds and streams in wooded areas. Lejogaster larvae have<br />
been found in <strong>the</strong> decaying vegetation <strong>of</strong> a floating mat <strong>of</strong> Typha and o<strong>the</strong>r plants in a<br />
pond. The posterior spiracles <strong>of</strong> <strong>the</strong>se two genera are not modified in<strong>to</strong> piercing sty lets.<br />
Larvae <strong>of</strong> Brachyopa live in sap runs or under dead bark, those <strong>of</strong> Sphegina in<br />
decaying wood and those <strong>of</strong> <strong>the</strong> rare Hammerschmidtia probably in dead wood <strong>of</strong> birch<br />
and aspen around which <strong>the</strong> adults have been found.<br />
Lundbeck (1916) found larvae and pupae <strong>of</strong> Neoascia geniculata (Meigen) in flood<br />
refuse and it is assumed that <strong>the</strong>y probably occur in marshy soils.<br />
Callicerini. Coe (1938, 1939) found larvae <strong>of</strong> Callicera (figs 265-268) in a wet decaying<br />
cavity in <strong>the</strong> fork <strong>of</strong> a pine, some <strong>of</strong> which <strong>to</strong>ok five years <strong>to</strong> pupate. The posterior<br />
spiracles have straight slits and <strong>the</strong> anal papillae are very long. It is assumed that <strong>the</strong><br />
o<strong>the</strong>r two members <strong>of</strong> <strong>the</strong> genus have similar life-his<strong>to</strong>ries; all are rare.<br />
Pelecocerini. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> our only species, Pelecocera tricincta Meigen,<br />
are unknown.<br />
Eumerini. The larvae <strong>of</strong> Eumerus tuberculatus Rondani and E. strigatus (Fallen) are<br />
well known pests <strong>of</strong> Narcissus bulbs and are known as small narcissus flies or lesser bulb<br />
flies (<strong>the</strong> large bulb-fly being Merodon). E. strigatus larvae (figs 262-264) also attack<br />
Iris, parsnip roots, onions and on <strong>the</strong> continent have attacked pota<strong>to</strong>es. The two<br />
species are not easily distinguished in <strong>the</strong> larval <strong>stages</strong> (Hodson, 1932a). The <strong>immature</strong><br />
<strong>stages</strong> <strong>of</strong> <strong>the</strong> remaining two <strong>British</strong> species are unknown but presumably have similar<br />
habits and could well repay investigation. Larvae <strong>of</strong> o<strong>the</strong>r families(e.g. Scathophagidae)<br />
also occur in Narcissus bulbs (see index).<br />
Microdontini. The remarkable larvae <strong>of</strong> Microdon (figs 269- 270) live in ants' nests<br />
where <strong>the</strong>y feed on <strong>the</strong> minute pellets <strong>of</strong> food discarded by <strong>the</strong> ants. The biology <strong>of</strong> M .<br />
mutabilis (L.) is described by Donisthorpe (1927) and that <strong>of</strong> M. eggeri Mik by Syms<br />
( 1935).<br />
<strong>Vol</strong>ucellini. <strong>Vol</strong>ucella larvae (figs 271- 276) are scavengers in <strong>the</strong> nests <strong>of</strong> bumble bees<br />
(Bombus), social wasps (Vespula) and <strong>the</strong> hornet (Vespa). When wasps nest in ro<strong>of</strong><br />
spaces it is not uncommon for <strong>Vol</strong>ucella larvae (usually V. inanis L., which lacks <strong>the</strong><br />
long anal papillae <strong>of</strong> o<strong>the</strong>r species, fig. 271) <strong>to</strong> leave <strong>the</strong> nests and appear (via light<br />
83
fittings) in <strong>the</strong> upstairs rooms <strong>of</strong> houses whence <strong>the</strong>y may be sent in <strong>to</strong> en<strong>to</strong>mologists for<br />
identification by environmental health <strong>of</strong>ficers. I have seen larvae <strong>of</strong> V. pellucens, which<br />
had obviously dropped from such a situation, served in a restaurant meal!<br />
The classic and beautiful work on <strong>Vol</strong>ucella is by Kiinckel d'Herculais ( 1875- 1881)<br />
but an illustrated key <strong>to</strong> species is provided by Hartley ( 1961 ).<br />
Sericomyiini. Sericomyia species have larvae <strong>of</strong> <strong>the</strong> 'rat-tailed maggot' type similar <strong>to</strong><br />
those <strong>of</strong>Eristalini. S.lappona (L.) has been found in disturbed peat lying in water at <strong>the</strong><br />
end <strong>of</strong> a derelict cutting (Hartley, 1961).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Arc<strong>to</strong>philafulva (Harris) are unknown, but a female has been<br />
seen ovipositing in deep, water-filled horse ho<strong>of</strong> prints along a shaded muddy path near<br />
a stream (Stubbs & Falk, 1983).<br />
Xylotini. Xylota larvae resemble those <strong>of</strong> Syritta (see below) and have been found in<br />
very wet decomposing silage, wet sawdust <strong>of</strong> an old saw pit, exudate and rot-holes <strong>of</strong><br />
yew (Taxus baccata L.), and wet decaying beech stumps (Fagus sylvatica L.).<br />
Larvae <strong>of</strong> Xylo<strong>to</strong>mima nemorum (F.) have been found overwintering under bark <strong>of</strong><br />
recently felled logs and branches lying in wet situations.<br />
The short-tailed larvae <strong>of</strong> a non-<strong>British</strong> species <strong>of</strong> Brachypalpus has been found in <strong>the</strong><br />
decaying wood <strong>of</strong> willow trees (on <strong>the</strong> continent).<br />
The very rare Caliprobola speciosa (Rossi) has been reared from a rotten beech<br />
stump.<br />
Syritta pipiens (L.) (figs 277- 278) breeds in various kinds <strong>of</strong> decaying organic matter<br />
such as manure heaps and <strong>the</strong> edge <strong>of</strong> silage clamps where <strong>the</strong>y may occur in very large<br />
numbers. They may also be found as saprophages in rotting bulbs (<strong>the</strong>y do not attack<br />
healthy bulbs) where <strong>the</strong>y could be confused with Eumerus larvae (see above).<br />
The puparium <strong>of</strong> Tropidia se ita (Harris) was found among flood refuse by Lundbeck<br />
( 1916) and an American species has been reared from human excrement near <strong>the</strong> mouth<br />
<strong>of</strong> a sewer and also in masses <strong>of</strong> rotting pota<strong>to</strong>es.<br />
Pocota has a short-tailed larva which has been found in rot-holes high above <strong>the</strong><br />
ground in various trees (Aubertin, 1928). Criorhina berberina (F.) has been reared from<br />
<strong>the</strong> slimy decay at <strong>the</strong> base <strong>of</strong> a fallen birch (Betula) trunk. The larva appears <strong>to</strong> be <strong>of</strong><br />
<strong>the</strong> short-tailed type, whereas <strong>the</strong> larvae <strong>of</strong> some North American species are rat-tailed.<br />
The early <strong>stages</strong> <strong>of</strong> Blerafallax (L.) are unknown but females have been seen laying<br />
eggs in sap exuding from beech and oak trees.<br />
Merodontini. Merodon equestris (F.) is <strong>the</strong> large bulb fly (or Narcissus fly), <strong>the</strong> larvae<br />
(figs 279- 280) <strong>of</strong> which attack narcissi and various o<strong>the</strong>r bulbs and can be a major pest<br />
<strong>to</strong> growers (see also Eumerus, <strong>the</strong> lesser bulb flies, above). Larvae have also been found<br />
in bulbs <strong>of</strong> <strong>the</strong> blue-bell (Endymion nonscriptus (L.)). Hod son ( 1932b) gives an account<br />
<strong>of</strong> <strong>the</strong> biology. Eggs (fig. <strong>10</strong>12) are laid on <strong>the</strong> neck <strong>of</strong> <strong>the</strong> bulb or on its dying leaves.<br />
Eristalini. The genera <strong>of</strong> this tribe have larvae <strong>of</strong> <strong>the</strong> rat-tailed maggot type (fig. 281 ),<br />
which may be separated by <strong>the</strong> following simplified key (Hartley, 1961 and Dolezil,<br />
1972 provide keys <strong>to</strong> some species):<br />
Pubescence absent on dorsum <strong>of</strong> last six abdominal segments. <strong>An</strong>terior spiracles especially<br />
long and pointed. Exclusively in tree rot-holes high above <strong>the</strong> ground and rarely seen until<br />
trees are felled . . . . . . . . . . . . . . . . . . . M a/Iota<br />
Pubescence usually present on dorsum <strong>of</strong> abdominal segments. <strong>An</strong>terior spiracles shorter.<br />
Larvae usually found in aquatic or semi-aquatic situations with high organic content<br />
occasionally in more accessible tree rot-holes . . . . . . . . . . . 2<br />
2 Abdominal prolegs with s<strong>to</strong>ut well sclerotized crochets in two distinct semicircular rows;<br />
anterior spiracle pale brownish . . . . . . . . . . Myathropa<br />
84
Abdominal pro legs with crochets in more than two distinct rows which are not semicircular<br />
on first three abdominal segments; if crochets tend <strong>to</strong> be in two rows <strong>the</strong>n <strong>the</strong> anterior<br />
spiracles are dark brown . . . . . . . . . . . . . . . . . . 3<br />
3 Tracheal trunks undulating (as seen through cuticle) . . Helophilus<br />
Tracheal trunks straight . . . Parhelophilus, <strong>An</strong>asimyia, Lejops, Eristalis, Eristalinus<br />
Mallota cimciciformis (Fallen) is ano<strong>the</strong>r tree hole breeder, usually in elm or horsechestnut,<br />
usually at a considerable height from <strong>the</strong> ground, and <strong>the</strong>refore rarely<br />
encountered until trees are felled .<br />
Larvae <strong>of</strong> Myathropajiorea (L.) are found in rot holes and leaf-filled water pockets<br />
in trees, especially beech, and also in wooden water butts with an accumulation <strong>of</strong><br />
decaying leaves at <strong>the</strong> bot<strong>to</strong>m.<br />
Helophilus pendulus (L.) usually breeds in farmyard drains, very wet manure,<br />
decomposing silage etc. (sometimes in company with Eristalis). H. hybridus Loew and<br />
species <strong>of</strong> Parhelophilus and <strong>An</strong>asimyia occur in ponds with a dense stand <strong>of</strong> Typha<br />
(reed mace). The larva <strong>of</strong> <strong>the</strong> rare Lejops vittata (Meigen) is unknown but should be<br />
sought in coastal marshes with growth <strong>of</strong> Scirpus maritimus among which adults have<br />
been found.<br />
Eristalis tenax (L.) breeds in places with a high animal sewage content such as<br />
farmyard drains. It also breeds in <strong>the</strong> liquid putrefaction from carcases and (because <strong>of</strong><br />
<strong>the</strong> bee-like adult) is <strong>the</strong> so called oxen-born bee <strong>of</strong> <strong>the</strong> ancients, which is shown coming<br />
from a lion carcase on <strong>the</strong> tin <strong>of</strong> a well known brand <strong>of</strong> golden syrup, accompanied by<br />
<strong>the</strong> quotation (from Samson) 'Out <strong>of</strong> <strong>the</strong> strong came forth sweetness' (see also Medical<br />
& Veterinary section). E. pertinax (Scopoli), E. nemorum (L.), E. arbus<strong>to</strong>rum (L.) and<br />
E. intricarius(L.) breed in organically polluted (cattle droppings, etc.) drains and pools.<br />
Larvae <strong>of</strong> E. abusivus Coli in have been found in mud at <strong>the</strong> edge <strong>of</strong> a moorland pond.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> o<strong>the</strong>r species require investigation.<br />
Eristalinus sepulchralis (L.) occurs in ponds with a large amount <strong>of</strong> decayed<br />
vegetation, but usually with little sewage contamination. E. aeneus (Scopoli) occurs in<br />
seashore pools containing rotting seaweed.<br />
Series Schizophora<br />
Acalyptratae<br />
Conopidae<br />
(Figs: larvae 282- 292, eggs <strong>10</strong>13- <strong>10</strong>16, pupa 1165)<br />
About 800 species <strong>of</strong> Conopidae in 47 genera have been described throughout <strong>the</strong><br />
world and 25 species representing 7 genera in 2 sub-families occur in Britain. The<br />
<strong>British</strong> representatives <strong>of</strong> <strong>the</strong> family are internal parasites <strong>of</strong> adult bees and wasps. The<br />
<strong>immature</strong> <strong>stages</strong> are poorly known and only 4 <strong>British</strong> species in 4 genera have been<br />
described in <strong>the</strong> larval stage. The biology and known host records for world species are<br />
reviewed by Smith (1966) and o<strong>the</strong>r papers are cited below. Smith & Peterson (1987)<br />
provide keys <strong>to</strong> genera for <strong>the</strong> eggs and larvae so far described.<br />
Conopinae. Larval mouthparts and posterior spiracles <strong>of</strong> Conops vesicularis L. ,<br />
recovered from <strong>the</strong> puparium, are described by Dusek (1964) (fig. 285). This species<br />
parasitises Bombus muscorum (L.). Host records for o<strong>the</strong>r <strong>British</strong> Conops species are<br />
C.jiavipes L. on Bombus lapidarius (L.) and Osmia, and C. quadrifasciatus De Geer also<br />
on Bombus lapidarius. The hosts <strong>of</strong> C. strigata Wiedemann and C. ceriaeformis Meigen<br />
are unknown.<br />
85
The hosts <strong>of</strong> <strong>the</strong> two <strong>British</strong> species <strong>of</strong> Leopoldius are unknown but <strong>the</strong> adults <strong>of</strong>ten<br />
occur on ivy blossom among wasps ( Vespula) and on <strong>the</strong> continent L. coronatus<br />
Rondani has been seen ovipositing on and has been reared from Vespula germanica<br />
(F.).<br />
Physocephala rufipes (F.) is parasitic on Bombus species. De Meijere (1912b) reared<br />
this species from Bombus pascuorum (Scopoli) ( = agrorum (F.)) and B. lapidarius and<br />
described <strong>the</strong> <strong>immature</strong> <strong>stages</strong>. Cumber (1949) reared it from six species <strong>of</strong> Bombus in<br />
<strong>the</strong> London area and found that up <strong>to</strong> 13% <strong>of</strong> individuals <strong>of</strong> some species were<br />
parasitized. P. nigra (De Geer) is parasitic on Bombus muscorum.<br />
Myopinae. De Meijere ( 1912b) described <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Zodion cinereum (F.)<br />
(figs 286--287) which he reared from Hylaeus quadricinctus (F.), Ha/ictus rubicundus<br />
Christ and Lasioglossum nigripes Lepeletier (=H. nylanderi Perez). In Russia <strong>the</strong><br />
species has been recorded from <strong>the</strong> honey-bee, Apis mellifera L. (Zimina, 1968).<br />
The larva <strong>of</strong> Myopa is unknown and <strong>the</strong> hosts <strong>of</strong> <strong>the</strong> <strong>10</strong> <strong>British</strong> species are<br />
unrecorded. However, in <strong>the</strong> <strong>British</strong> Museum (Natural His<strong>to</strong>ry) collection <strong>the</strong>re is a<br />
specimen <strong>of</strong> M . fasciata Miegen labelled 'Host: <strong>An</strong>drena cetii Schrank = marginata<br />
Fab.' 'England: Devon, Bovey Tracey, ix.l9, R. C. L. Perkins (BM 1942-95)'. On<br />
<strong>the</strong> continent unidentified or non-<strong>British</strong> members <strong>of</strong> <strong>the</strong> genus have been reared or<br />
associated with species <strong>of</strong> Eucera, Bombus, <strong>An</strong>drena, Colletes and Vespula.<br />
The larva <strong>of</strong> Thecophora (figs 289- 292) has been described only for a Nearctic species<br />
(Smith, 1966) associated with various Ha/ictus and Lasioglossum species. In Britain<br />
T. atra (F.) and T. fulvipes Robineau-Desvoidy, and on <strong>the</strong> continent T. pusilla<br />
(Meigen), have all been found among Ha/ictus colonies. In Russia two (non-<strong>British</strong>)<br />
species are recorded from <strong>the</strong> honey-bee, Apis mellifera (Zimina, 1968).<br />
De Meijere (1912b) described <strong>the</strong> larva <strong>of</strong> Sicusferrugineus (L.) (fig. 288) which has<br />
been recorded from seven species <strong>of</strong> Bombus on <strong>the</strong> continent.<br />
Tephritidae<br />
(Figs: larvae 293-314, eggs <strong>10</strong>17- <strong>10</strong>20, puparia 1166--1168,<br />
mines 1293- 1295, galls 1316--1317)<br />
The Tephritidae ( = Trypetidae) (fruit flies) contains some 4,500 species and <strong>of</strong> all <strong>the</strong><br />
families <strong>of</strong>Diptera is said <strong>to</strong> cause <strong>the</strong> most economic damage <strong>to</strong> plants.<br />
The larval habits <strong>of</strong>Tephritidae fall in<strong>to</strong> four major groups as follows:<br />
I. Developing in flower heads (mostly Compositae-thistles, sow-thistles, hawkweeds, etc.),<br />
e.g. Urophora, Myopites, Xyphosia, Trupanea, Ensina, Noeeta, Tephritis, etc.<br />
2. Leaf-miners, e.g. Euleia, Trypeta, Vida/ia, Myoleja.<br />
3. Developing in stems or roots, e.g. Oxyna, Campiglossa, Cerajocera, Dithryca, Platyparea,<br />
Platyparella.<br />
4. Developing in fruits, e.g. Rhagoletis, <strong>An</strong>omoia, Ceratitis, etc.<br />
Some <strong>of</strong> <strong>the</strong> non-<strong>British</strong> pest species are included below as <strong>the</strong>y are frequently found<br />
in imported fruit and sent <strong>to</strong> en<strong>to</strong>mologists for identification. There are 76 native<br />
<strong>British</strong> species representing 33 genera and 3 subfamilies.<br />
A useful summary <strong>of</strong> <strong>the</strong> biology <strong>of</strong> fruit flies is provided by Christenson & Foote<br />
(1960). Phillips (1946), Varley (1937), Persson (1963), Dirlbeck & Dirlbeck (1962),<br />
Kandybina (1977) and G. H. Berg (1979) are useful for <strong>the</strong> identification <strong>of</strong> some<br />
Tephritid larvae that occur in Britain and fur<strong>the</strong>r information, full host plant lists and<br />
some keys (<strong>to</strong> puparia) are provided in <strong>the</strong> <strong>British</strong> Handbook <strong>to</strong> <strong>the</strong> family (White,<br />
1988). Although <strong>the</strong> host plants <strong>of</strong> most <strong>British</strong> species are known <strong>the</strong> <strong>immature</strong> <strong>stages</strong><br />
<strong>of</strong> many are still inadequately described or unknown. Since White (1988) gives such a<br />
detailed treatment <strong>of</strong> <strong>the</strong> family my comments on host-plants and biology are restricted<br />
86
<strong>to</strong> <strong>the</strong> more frequently encountered genera and species, including those <strong>of</strong> economic<br />
importance.<br />
Trypetinae<br />
Ceratitini. Ceratitis capitata (Wiedemann), <strong>the</strong> well-known Mediterranean fruit fly,<br />
is not a native <strong>British</strong> species but its larvae (figs 293- 296) occur in cherries, plums,<br />
peaches, apricots, pears, oranges and <strong>to</strong>ma<strong>to</strong>es imported in<strong>to</strong> this country. O<strong>the</strong>r<br />
non-<strong>British</strong> Tephritidae, e.g. Dacus, <strong>An</strong>estrepha, occurring in imported fruits may be<br />
identified with <strong>the</strong> work <strong>of</strong> Berg (1979) and <strong>British</strong> records are discussed by White<br />
( 1988) (see also Rhagoletis cerasi (L.) below).<br />
Trypetini. <strong>An</strong>omoia ( = Phagocarpus) purmunda (Harris) larvae are found in <strong>the</strong> fruits<br />
('haws') <strong>of</strong> hawthorn but also occur in <strong>the</strong> berries <strong>of</strong> Berberis, Pyracantha and<br />
Co<strong>to</strong>neaster. Platyparella discoidea (F.) is said <strong>to</strong> occur in <strong>the</strong> stems <strong>of</strong> Campanula<br />
latifolia L. Platyparea poeciloptera (Schrank) is <strong>the</strong> horticulturalist's asparagus fly, <strong>the</strong><br />
larvae (figs 297- 299) <strong>of</strong> which damage <strong>the</strong> stems <strong>of</strong> asparagus (Lesne, 1913) (fig. 1293)<br />
though no attacks have been reported in Britain since <strong>the</strong> 1930s. Euleia heracleii (L.) is<br />
<strong>the</strong> celery fly, <strong>of</strong> economic importance <strong>to</strong> horticulturalists, <strong>the</strong> larvae <strong>of</strong> which, in<br />
addition <strong>to</strong> celery (Apium graveolens L.) also mine <strong>the</strong> leaves <strong>of</strong> parsnip (Peucedanum<br />
sativum Bentham & Hooker), Heracleum sphondylium L., Sium latifolium L., <strong>An</strong>gelica<br />
sylvestris L. and o<strong>the</strong>r Umbelliferae. Acidia cognata (Wiedemann) larvae mine <strong>the</strong><br />
leaves <strong>of</strong> Tussilago far far aL. and Petasites ovatus Hill.<br />
Trypeta and Vidalia species mine leaves <strong>of</strong> Artemisia, Eupa<strong>to</strong>rium , Senecio, etc.<br />
Rhagoletis alternata (F.) larvae occur in <strong>the</strong> fruits ('hips') <strong>of</strong> wild and cultivated roses.<br />
R. cerasi (L.) is <strong>the</strong> European cherry fly and its larvae (figs 304-306) are frequently<br />
found in imported cherries. Larvae <strong>of</strong> Gonioglossum wiedemanni (Meigen) are found in<br />
<strong>the</strong> berries <strong>of</strong> Bryonia dioica.<br />
Euphrantini. The larva <strong>of</strong> Euphranta <strong>to</strong>xoneura (Loew) is one <strong>of</strong> <strong>the</strong> very few<br />
tephritids that departs from <strong>the</strong> phy<strong>to</strong>phagous habit and lives as a preda<strong>to</strong>r/brood<br />
parasite in sawfly galls <strong>of</strong> <strong>the</strong> genus Pontania (Kopelke, 1984, 1985).<br />
Terelliini. Larvae <strong>of</strong> Chae<strong>to</strong>s<strong>to</strong>mella cylindrica (Robineau-Desvoidy) occur in <strong>the</strong><br />
flower-heads <strong>of</strong> Centaurea. Chae<strong>to</strong>rellia and Cerajocera larvae (figs 311-312) are found<br />
in <strong>the</strong> flower-heads <strong>of</strong> Centaurea and <strong>the</strong> latter also in Arctium. The larva <strong>of</strong> Cerajocera<br />
cera<strong>to</strong>cera Hendel has a very characteristic anal plate (fig. 312).<br />
Larvae <strong>of</strong> Orellia falcata (Scopoli) are found in <strong>the</strong> root-s<strong>to</strong>ck <strong>of</strong> Tragopogon<br />
pratense L.<br />
Terellia species occur in flower-heads as follows: T. colon (Meigen) in Centaurea<br />
scabiosa L.; T. ruficauda (F.) in Cirsium; T. vectensis Collin on Serratula tinc<strong>to</strong>ria L.;<br />
T. win<strong>the</strong>mi (Meigen) on Carduus acanthoides L.<br />
Xyphosia miliaria (Schrank) occurs in flower-heads <strong>of</strong> Cirsium .<br />
Myopitinae<br />
The larvae <strong>of</strong> Urophorajaceana (Hering), <strong>the</strong> knapweed gall fly (figs 307- 3<strong>10</strong>), form<br />
galls (fig. 1317) in <strong>the</strong> flower-heads <strong>of</strong> Centaurea nigra L. and this was <strong>the</strong> subject <strong>of</strong> a<br />
classical ecological study by Varley (1947). For many years U.jaceana was erroneously<br />
recorded as U. solstitialis (L.) which inhabits galls on Carduus nutans L. U. cuspidata<br />
(Meigen) and U. quadrifasciata (Meigen) also occur in <strong>the</strong> flower-heads <strong>of</strong> Centaurea.<br />
U. spoliata (Haliday) lives in <strong>the</strong> flower-heads <strong>of</strong> Serratula tinc<strong>to</strong>ria, U. stylata (F.) in<br />
galls in flower-heads <strong>of</strong> Cirsium vulgare (Save) Ten. and U. car dui (L.) forms galls on <strong>the</strong><br />
stems <strong>of</strong> Cirsium.<br />
87
Our two Myopites species form hard galls in <strong>the</strong> flower-heads <strong>of</strong> Pulicaria dysenterica<br />
L. and !nu/a crithmoides L. Ensina sonchi (L.) occurs in <strong>the</strong> flower-heads <strong>of</strong> a range <strong>of</strong><br />
plants including Hypochaeris , Sonchus, Picris, Leon<strong>to</strong>don, Tragopogon, Aster, etc.<br />
Tephritinae<br />
Dithrycini. The larva <strong>of</strong> Dythryca guttularis (Meigen) forms a gall at <strong>the</strong> stem base <strong>of</strong><br />
Achillea millefolium L. Noeeta pup illata (Fallen) is found in <strong>the</strong> swollen flower-heads <strong>of</strong><br />
Hieracium spp.<br />
Tephritini. lcterica westermanni (Meigen) occurs in <strong>the</strong> flower-heads <strong>of</strong> Senecio spp.<br />
The food-plant <strong>of</strong> Acinia corniculata (Zetterstedt) is probably Centaurea nigra (on <strong>the</strong><br />
continent it has been reared from <strong>the</strong> related C.jacea).<br />
Oxyna jlavipennis (Loew) larvae live in fleshy galls on <strong>the</strong> roots <strong>of</strong> Achillea<br />
millefolium L. and those <strong>of</strong> 0. nebulosa (Wiedemann) in galls on <strong>the</strong> roots<strong>to</strong>ck <strong>of</strong><br />
Chrysan<strong>the</strong>mum leucan<strong>the</strong>mum L. 0. parietina (L.) larvae live in galls on <strong>the</strong> stems <strong>of</strong><br />
Artemisia spp.<br />
Campiglossa grandinata (Rondani) lives in a gall on <strong>the</strong> stems <strong>of</strong> Solidago virgaurea<br />
L. and C. argyrocephala (Loew) forms a rosette gall in <strong>the</strong> flowerhead <strong>of</strong> Achillea<br />
ptarmica.<br />
Sphenella marginata (Fallen) is found in <strong>the</strong> swollen flower-heads <strong>of</strong> Senecio species.<br />
Paroxyna are mostly found in flower-heads: P. absinthii (F.) on Artemisia; P.<br />
loewiana Hen del on Solidago virgaurea (L.); P. plantaginis (Haliday) on Aster tripolium<br />
L.; P. product a (Loew) on Sonchus, Hypochoeris, Crepis, Taraxacum , Leon<strong>to</strong>don, etc.<br />
Larvae <strong>of</strong> Tephritis species occur in flower-heads <strong>of</strong>Compositae.<br />
Trupanea amoena (Frauenfeld) occurs in flower-heads <strong>of</strong> Lactuca, Picris and<br />
Sonchus. T. stellata (Fuessly) is found in flower-heads <strong>of</strong> <strong>An</strong><strong>the</strong>mis, Artemisia, Aster,<br />
Centaurea, Crepis, Hieracium, Matricaria and Senecio.<br />
Acanthiophilus helianthi (Rossi) occurs in flower-heads <strong>of</strong> Centaurea.<br />
Platys<strong>to</strong>matidae<br />
(Figs: larvae 315- 321)<br />
This is one <strong>of</strong> <strong>the</strong> largest families <strong>of</strong> Acalyptratae with over I ,000 species <strong>of</strong> which<br />
about 60 occur in <strong>the</strong> Palaearctic region but only 2 species representing 2 genera occur<br />
in <strong>the</strong> U.K., nei<strong>the</strong>r <strong>of</strong> which is known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Platys<strong>to</strong>ma seminationis (L.) has been reared from a fungus (Tricholomopsis rutilans<br />
Quelet). The larva <strong>of</strong> <strong>the</strong> European P. lugubre (Robineau-Desvoidy) was briefly<br />
described by Perris (1856) who found it on soil beneath a piece <strong>of</strong> dead wood (figs<br />
315- 316). Hennig (1945) described larval details from a puparium <strong>of</strong> P. euphorbiinum<br />
Enderlein from <strong>the</strong> Canary Isles (figs 317- 319).<br />
Larvae <strong>of</strong> Rivellia species (figs 320-321) have been found feeding on <strong>the</strong> root nodules<br />
<strong>of</strong> various leguminous plants in Zaire (Seeger & Maldague, 1960) and o<strong>the</strong>r countries,<br />
but <strong>the</strong> life-his<strong>to</strong>ry <strong>of</strong> <strong>the</strong> only <strong>British</strong> species, R. syngenesiae (F.), is unknown.<br />
Otitidae<br />
(Figs: larvae 322- 337, eggs <strong>10</strong>21-<strong>10</strong>22, puparia 1169)<br />
Over 500 species <strong>of</strong> Otitidae have been described throughout <strong>the</strong> world. The little<br />
that is known <strong>of</strong> <strong>the</strong>ir biology is summarised by Alien & Foote (1967). The <strong>immature</strong><br />
88
<strong>stages</strong> have been found in decaying or damaged vegetable substances and in animal<br />
faeces. Twenty-one species in 11 genera representing two subfamilies occur in Britain.<br />
Ulidiinae. The larvae <strong>of</strong> Physiphora demandata (F.) (figs 322-324) have been found<br />
in corn silage, clover fodder, rotting narcissus bulbs and dung (Alien, E. J. & Foote,<br />
1967). The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Ulidia erythrophthalma Meigen and our two Homalocephala<br />
species are undescribed but puparia <strong>of</strong> H. albitarsis Zetterstedt have been<br />
found under <strong>the</strong> bark <strong>of</strong> dead and dying coniferous trees in Sweden.<br />
Otitinae. The larva <strong>of</strong> Myennis oc<strong>to</strong>punctata (Coquebert) (figs 325- 328) was<br />
described by V os de Wilde ( 1935) from larvae found under <strong>the</strong> bark <strong>of</strong> dead trees.<br />
Seioptera vibrans (L.) has been reared from manure, decaying fruit, vegetables,<br />
narcissus bulbs, onions and dung. The larvae (figs 329- 331) have been described by<br />
Lobanov (1958) and Alien & Foote (1967). The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Herina (7 <strong>British</strong><br />
species) are unknown.<br />
The larva <strong>of</strong> Ceroxys urticae (L.) (figs 332-335) has been found in manure and<br />
decaying vegetation and is described by Lobanov (1964).<br />
Melieria species are unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>, as is <strong>the</strong> larva <strong>of</strong> Tetanops<br />
myopinus Fallen. The adult <strong>of</strong>T. myopinus is rarely seen in this country but occurs in<br />
coastal areas and has been seen resting on marram grass (Smith, 1955b). In North<br />
America T. myopaef ormis (Roder) (figs 336--337) is an important pest <strong>of</strong> sugarbeet<br />
(Gojmerac, 1956) and T. luridipennis Loew has been reared from decaying narcissus<br />
bulbs. Lobanov ( 1972) has described <strong>the</strong> larva <strong>of</strong>T. sintenisi Beck er.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Otites guttata (Meigen) and Dorycera graminum (F.)<br />
are undescribed but <strong>the</strong> latter is said <strong>to</strong> oviposit on <strong>the</strong> 'ovaries' <strong>of</strong> Tamus (Seguy,<br />
1934).<br />
Micropezidae<br />
(Figs: larvae 338- 352, puparia 1170-1171)<br />
The Micropezidae contains some 450 species <strong>of</strong> which 9 species representing 3 genera<br />
in 3 subfamilies occur in Britain. The larvae are saprophagous in decaying vegetation<br />
and some are phy<strong>to</strong>phagous on <strong>the</strong> root nodules <strong>of</strong> leguminous and o<strong>the</strong>r plants.<br />
Micropezinae. The larva <strong>of</strong> Micropeza corrigiolata (L.) (figs 338-342) has been<br />
described by M tiller (1957). They bore in<strong>to</strong> <strong>the</strong> root nodules <strong>of</strong> leguminous crops, e.g.<br />
field pea (Pisum arvense L.), red clover (Trifolium pratense L.) and alfalfa (Medicago<br />
sativa L.). The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> M. lateralis Meigen are unknown.<br />
Taeniapterinae. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> only <strong>British</strong> species, Rainieria calceata<br />
(Fallen), are unknown but larval details from a puparium <strong>of</strong> <strong>the</strong> Nearctic R. antennaepes<br />
(Say) found in <strong>the</strong> crotch <strong>of</strong> a large American elm have been described (Steyskal, 1964)<br />
and are illustrated here (figs 343- 345).<br />
Calobatinae. The larvae <strong>of</strong> Calobata cibaria (L.) (figs 346--347) and C. petronella (L.)<br />
(fig. 351) have been described by Brindle (1965a) from specimens found in a well<br />
decayed grass heap. Lobanov (1960) has also described <strong>the</strong> larva <strong>of</strong> C. petronella<br />
from 'refuses and strongly mineralized aggregations <strong>of</strong> swine manure' from which<br />
<strong>the</strong> third instar migrates in<strong>to</strong> drier adjacent soil areas with decreased organic content<br />
for <strong>the</strong> pupose <strong>of</strong> hibernation. <strong>An</strong> interesting feature <strong>of</strong> <strong>the</strong> third stage larva <strong>of</strong> <strong>the</strong><br />
Nearctic species C. vittata (Walker) is <strong>the</strong> presence <strong>of</strong> 4 slits in <strong>the</strong> posterior spiracle<br />
(fig. 352). The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> remaining 4 <strong>British</strong> species <strong>of</strong> <strong>the</strong> genus are<br />
unknown.<br />
89
Megamerinidae<br />
(Figs: larvae 353- 354, puparium 1172)<br />
Only 13 species <strong>of</strong> this family are known, 11 in <strong>the</strong> Oriental region and 2 Palaearctic<br />
species <strong>of</strong> which Megamerina dolium (F.) occurs in Britain.<br />
The larvae live under <strong>the</strong> bark <strong>of</strong> dead and dying broad-leaved trees <strong>to</strong>ge<strong>the</strong>r with <strong>the</strong><br />
larvae <strong>of</strong> o<strong>the</strong>r xylophilous Diptera and are apparently predaceous or necrophagous.<br />
Hennig (1943b) has described <strong>the</strong> larva <strong>of</strong> M . dolium (figs 353- 354).<br />
Tanypezidae<br />
(Figs: larvae 355- 360)<br />
Twenty species <strong>of</strong>Tanypezidae have been described (mostly Neotropical) <strong>of</strong> which<br />
<strong>the</strong> only Palaearctic species, Tanypeza longimana Fallen, occurs in Britain. The larval<br />
<strong>stages</strong> (figs 355- 360) have been reared in <strong>the</strong> labora<strong>to</strong>ry on watermelon rind and pulp<br />
from eggs laid by a gravid female from a densely shaded colony <strong>of</strong> yellow skunk<br />
cabbage (Lysichitum americanum Hul<strong>to</strong>n & St. John, Araceae) in <strong>the</strong> U.S.A. (Foote,<br />
1970). None <strong>of</strong> <strong>the</strong> larvae formed puparia so it is not certain if <strong>the</strong>y are naturally<br />
saprophagous. The species is rare in Britain and has been found among streamside<br />
vegetation.<br />
Psilidae<br />
(Figs: larvae 361-372, egg <strong>10</strong>23, puparium 1173)<br />
There are about 170 species <strong>of</strong>Psilidae worldwide <strong>of</strong> which 55 are Palaearctic and 26<br />
<strong>of</strong> <strong>the</strong>se species, representing 3 genera, occur in Britain. The larvae are phy<strong>to</strong>phagous,<br />
usually boring in <strong>the</strong> roots <strong>of</strong> plants, and some are pests. Some species develop under<br />
<strong>the</strong> bark <strong>of</strong> trees.<br />
De Meijere (1947) described a larva found in <strong>the</strong> base <strong>of</strong> a stem <strong>of</strong> ]uncus as Loxocera<br />
albiseta (Schrank) (figs 361-363) but Brindle (1965d) suggests that this description<br />
could in fact refer <strong>to</strong> ano<strong>the</strong>r Loxocera species, possibly L. ichneumonea (L.) ( = aristata<br />
(Panzer)). It would be a useful exercise <strong>to</strong> rear any psilid larvae found in ]uncus <strong>to</strong> settle<br />
this point, and perhaps elucidate <strong>the</strong> life-his<strong>to</strong>ry <strong>of</strong> o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> genus which<br />
remain unknown.<br />
Psi/a rosae (F.) is <strong>the</strong> carrot fly (Ash by & Wright, 1946) which is a well-known pest in<br />
most fields, allotments and gardens where <strong>the</strong> larvae (figs 364-367) mine in<strong>to</strong> <strong>the</strong> tap<br />
roots <strong>of</strong> carrots and render <strong>the</strong>m unsaleable. Parsnips, turnips and <strong>the</strong> crowns and<br />
basal stems <strong>of</strong> celery are also attacked as well as wild Umbelliferae (e.g. <strong>An</strong>thriscus,<br />
Heracleum). In carrots P. rosae larvae are yellow but from o<strong>the</strong>r plants <strong>the</strong>y are white.<br />
Hard man et al. (1985) provide a bibliography <strong>of</strong> P. rosae. Psi/a nigricornis Meigen is <strong>the</strong><br />
chrysan<strong>the</strong>mum s<strong>to</strong>ol miner which can be a serious horticultural pest. Under glass<br />
P. nigricornis and P. rosae may attack lettuce following chrysan<strong>the</strong>mums and celery<br />
respectively. P. nigricornis may also attack outdoor crops <strong>of</strong> lettuce (Margaret E. John<br />
pers. comm.). Osborne (1961) differentiates <strong>the</strong> larvae and puparia <strong>of</strong> <strong>the</strong> two species<br />
and notes that P. bicolor Meigen and P.limbatella (Zetterstedt) have also been found <strong>to</strong><br />
attack chrysan<strong>the</strong>mums but unfortunately no larval descriptions <strong>of</strong> any <strong>of</strong> <strong>the</strong>se o<strong>the</strong>r<br />
species <strong>of</strong> Psi/a occurring in Britain are available. See also Napomyza (Agromyzidae).<br />
The larvae <strong>of</strong> Chyliza vittata Meigen (figs 368- 372) occur in <strong>the</strong> stems <strong>of</strong> orchids (V os<br />
de Wilde, 1935). O<strong>the</strong>r species <strong>of</strong> Chyliza have been reared from orchids but <strong>the</strong> larvae<br />
are undescribed, e.g. C. extenuatum (Rossi) from bulbs <strong>of</strong> broomrape (Orobanche) ;<br />
C. scutellata F. from stems <strong>of</strong> <strong>the</strong> bird's-nest orchid (Neottia nidus-avis (L.)) and<br />
apparently <strong>the</strong> larva also causes woody galls on <strong>the</strong> stems <strong>of</strong> Spiraea. Several species <strong>of</strong><br />
Chyliza have been reared from tree bark, sap or resin in North America and Winter<br />
90
( 1988) describes <strong>the</strong> larva <strong>of</strong> C. fuscipennis (Robineau-Desvoidy) from resinous<br />
wounds on conifers in Britain.<br />
Helcomyzidae<br />
(Figs: larvae 373-379, egg <strong>10</strong>24, puparia 1174)<br />
This is a very small family <strong>of</strong> 12 species found in <strong>the</strong> Holarctic region and sub<br />
<strong>An</strong>tarctica (Falkland Is., Tierra del Fuego, S. Chile and Argentina). The larvae develop<br />
in rotting sea weeds on <strong>the</strong> sea shore. Two species representing 2 genera occur in Britain.<br />
The larvae <strong>of</strong> Helcomyza ustulataCurtis (figs 373-375) live in small fairly dry seaweed<br />
(wrack) beds lying on <strong>the</strong> sand in <strong>the</strong> high tide position (Egglishaw, 1960b). Earlier<br />
descriptions <strong>of</strong> <strong>the</strong> supposed larva <strong>of</strong> this species were erroneous (see Egglishaw, 1960b;<br />
Smith, 1981 ).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Heterocheila buccata (Fallen) (figs 376-379) are described by<br />
Backlund ( 1945a) and Egglishaw (1960a). The latter gives a full account <strong>of</strong> <strong>the</strong> biology<br />
and ecology <strong>of</strong> <strong>the</strong> species. Larvae <strong>of</strong> H. buccata are found only among <strong>the</strong> deep<br />
stipes <strong>of</strong> <strong>the</strong> wrack string (Laminaria) which are <strong>of</strong>ten partly buried in damp sand.<br />
Apparently <strong>the</strong>y do not occur among Fucus or <strong>the</strong> larger wrack beds.<br />
Dryomyzidae<br />
(Figs: larvae 38(}.-384, eggs <strong>10</strong>25-<strong>10</strong>27, puparium 1175)<br />
This little-known family is confined <strong>to</strong> <strong>the</strong> Holarctic region and only 15 species have<br />
been described <strong>of</strong> which two, representing one genus, occur in Britain. The <strong>immature</strong><br />
<strong>stages</strong> are associated with putrefying matter including excrement, carrion and rotting<br />
fungi , usually in woodland. <strong>An</strong> exception <strong>to</strong> this is <strong>the</strong> remarkable North American<br />
species Oedoparena glauca (Coquillett), <strong>the</strong> larva <strong>of</strong> which is a preda<strong>to</strong>r <strong>of</strong> intertidal<br />
barnacles (Burger et al., 1980).<br />
The larva <strong>of</strong> Dryomyza ana/is Fallen (figs 38(}.-384) is described by Smith (1981)<br />
from a dead pheasant and <strong>the</strong> species has also been reared from dead shrews (Disney,<br />
1973) and a dead fox (Smith, 1975). J. K. Barnes (1984) also describes <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong> and biology in <strong>the</strong> U .S.A. where it has been successfully reared in <strong>the</strong> labora<strong>to</strong>ry<br />
on hamburger, dead annelids, molluscs, insects, vertebrates and rotting fungi.<br />
However larvae failed <strong>to</strong> reach maturity when fed on rotting grass, decayed pumpkin<br />
flesh, decaying lettuce or cow manure. The egg (figs <strong>10</strong>25- <strong>10</strong>26) bears dorsolateral<br />
flanges and is particularly well adapted <strong>to</strong> survive on <strong>the</strong> type <strong>of</strong> substrate chosen for<br />
oviposition.<br />
D.flaveola (F.) has been reared from human and cow dung but only <strong>the</strong> egg (fig.<br />
I 027) has been described (Hin<strong>to</strong>n, 1960, 1981 ). It has long anterior horns as do some<br />
Drosophilidae.<br />
Chamaemyiidae<br />
(Figs: larvae 385-395, eggs <strong>10</strong>29-<strong>10</strong>30, puparia 1176-1179)<br />
The Chamaemyiidae (silver flies) contain about 183 species with a cosmopolitan<br />
distribution. The larvae are important preda<strong>to</strong>rs <strong>of</strong> aphids and Coccoidea<br />
(Homoptera) including many <strong>of</strong> <strong>the</strong> pest species. Twenty-five species representing 4<br />
genera in 2 subfamilies occur in Britain.<br />
Chamaemyiinae. Larvae <strong>of</strong> Chamaemyia feed on coccids living on grasses<br />
(Gramineae=Poaceae). In Russia, Tanasijtshuk (1970) has reared C. geniculata<br />
(Zetterstedt) (fig. 385) from a puparium found in <strong>the</strong> ovisac <strong>of</strong> a scale insect on cereal<br />
grass roots, and C. juncorum (Fallen) (fig. 386) from larvae feeding on scale insects<br />
91
(Balanococcus sp.) on Festuca ovina L. (and possibly a Metadenopus sp. on F. ovina<br />
subsp. sulcata). Collin (1966) found C. polystigma (Meigen) <strong>to</strong> be predaceous upon<br />
Pseudococcus phalaridis Green. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> remaining <strong>British</strong> species <strong>of</strong><br />
<strong>the</strong> genus are unknown but those <strong>of</strong> C.jlavipalpis Haliday should be sought on sandy<br />
coasts where <strong>the</strong> adults occur.<br />
Acrome<strong>to</strong>pia wahlbergi (Zetterstedt) is unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> but is<br />
probably predaceous on coccids or aphids on grasses among which <strong>the</strong> adults are<br />
usually found.<br />
The larvae <strong>of</strong> Parochthiphila coronata (Loew) (figs 387-389) were found feeding on<br />
<strong>the</strong> scale insect Pseudococcus aberrans Goux in <strong>the</strong> leaf sheath <strong>of</strong> couch grass by<br />
Tanasijtshuk (1963) in Russia. The same author suggests that o<strong>the</strong>r members <strong>of</strong> <strong>the</strong><br />
genus should be sought among scale insects in cereal leaf-sheaths and describes a new<br />
Russian species from such a situation in reeds.<br />
Leucopinae. Tanasijtshuk (1959), from a study <strong>of</strong> Leucopis species in <strong>the</strong> Crimea,<br />
divides <strong>the</strong> genus in<strong>to</strong> three biological groups:<br />
I. The eggs are deposited in <strong>the</strong> colonies <strong>of</strong> different species <strong>of</strong> aphids living openly. The larvae<br />
crawl on twigs and leaves <strong>of</strong> <strong>the</strong> plant and feed on aphids. They pupate ei<strong>the</strong>r on <strong>the</strong> plant<br />
or in <strong>the</strong> soil and hibernate as puparia. The number <strong>of</strong> generations per year is never less<br />
than two. L. melanopus Tanasijtshuk (fig. 391) belongs <strong>to</strong> this group.<br />
2. The eggs are deposited inside aphid galls. Both <strong>the</strong> feeding <strong>of</strong>larvae on aphids, and pupation,<br />
take place inside <strong>the</strong> galls. The number <strong>of</strong> generations per year and hibernation habits<br />
unknown. L. palumbi Rondani (not <strong>British</strong>) belongs <strong>to</strong> this group.<br />
3. The eggs are deposited in <strong>the</strong> ovisacs <strong>of</strong> coccids. The larvae develop and pupate in <strong>the</strong><br />
ovisacs. There are two generations per year but hibernation habits are unknown. So far<br />
only <strong>the</strong> subgenus Leucopomyia is placed in this group.<br />
A review <strong>of</strong> <strong>the</strong> <strong>British</strong> species including <strong>the</strong>ir known biology and habits is given by<br />
Smith (1963).<br />
All <strong>the</strong> <strong>British</strong> species <strong>of</strong> Leucopis (s.s.) (fig. 390) have been reared as follows:<br />
Leucopis albipuncta Zetterstedt from a bright yellow larva found feeding on aphids on a<br />
poppy (specimen in BM(NH)); L. atritarsis Tanasijtshuk among aphids on nettles; L.<br />
griseola (Fallen) (fig. 392) from larvae feeding on <strong>the</strong> aphid Eriosoma ulmi (L.) on elm;<br />
L. melanopus Tanasijtshuk (fig. 391) feeding on Dactynotus sp. on Lactuca viminae, on<br />
Brachycaudus car dui (L.) on Carduus, Aphis laburni Kaltenbach on Genista hispanica L.,<br />
Myzus persicae Sulz on henbane and on aphids from sage (all in <strong>the</strong> Crimea);<br />
L. morgei Smith from larvae feeding on <strong>the</strong> aphid Myzus ornatus Laing on Cydonia<br />
speciosa Sweet; L. puncticornis Meigen feeding on Brachycaudus cardui on Cirsium<br />
vulgare, Microlophium evansii (Theobald) and Aphis urticata Gmelin on Urtica dioica L.<br />
Leucopis ( Neoleucopis) obscura Haliday feeds on Pineus ( Adelges) pini (Gmelin in<br />
Linnaeus) on Scots pine and <strong>the</strong> o<strong>the</strong>r two <strong>British</strong> members <strong>of</strong> this subgenus appear <strong>to</strong><br />
do <strong>the</strong> same.<br />
Leucopis (Leucopomyia) silesiaca Eggers has been reared from among Eriococcus sp.<br />
on grass and from a hibernating egg mass <strong>of</strong> Eriopeltis festucae (Fonscolombe).<br />
Tanasijtshuk (1965) describes <strong>the</strong> larva (figs 393- 395).<br />
Leucopis (Lipoleucopis) praecox de Meijere feeds on Pineus pinion pine trees.<br />
Lauxaniidae<br />
(Figs: larvae 396-405, egg <strong>10</strong>28, puparium 1180)<br />
About 1500 species <strong>of</strong>Lauxaniidae ( =Sapromyzidae) have ben described throughout<br />
<strong>the</strong> world. Miller (1977) estimates that <strong>of</strong> <strong>the</strong>se about 5% have been reared but <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> <strong>of</strong> only about I 0 species have been described. The larval <strong>stages</strong> appear<br />
<strong>to</strong> be mostly saprophagous being found in fallen leaves, straw, rotting wood and birds'<br />
92
nests but a few are phy<strong>to</strong>phagous in root collars, stems, leaves <strong>of</strong> clover, <strong>the</strong> ovaries <strong>of</strong><br />
Viola and <strong>the</strong> leaf-like phylloclades <strong>of</strong> Opuntia cacti. No species appears <strong>to</strong> have any<br />
economic importance which probably explains <strong>the</strong> neglect <strong>of</strong> so large a family. I have<br />
seen probable lauxaniid larvae from rotting tissue at <strong>the</strong> base <strong>of</strong> winter barley stems<br />
and from <strong>the</strong> petiole <strong>of</strong> oil-seed rape but <strong>the</strong>se appeared <strong>to</strong> be saprophagous (J. A.<br />
Whiteway, pers. comm.). In Britain 46 species occur representing 11 genera. The<br />
<strong>immature</strong> <strong>stages</strong> and biology are reviewed by Miller & Foote (1975, 1976), and Miller<br />
(1977).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Trigonome<strong>to</strong>pusfrontalis (Meigen) are unknown but may be<br />
associated with reeds (Phragmites) among which <strong>the</strong> adults occur.<br />
Minettia inusta (Meigen) has been reared from <strong>the</strong> nest <strong>of</strong> a mole; M. lupulina (F.)<br />
from mines in leaflitter (Acer, Prunus, Alnus) and M. plumicornis (Fallen) from rotting<br />
leaves. The remaining seven <strong>British</strong> species have not been reared and only Minettia<br />
lupulina (Holarctic) has been described in <strong>the</strong> <strong>immature</strong> <strong>stages</strong> (Miller & Foote, 1976)<br />
and is illustrated here (figs 401-402).<br />
Sapromyza apicalis (Robineau-Desvoidy) has been reared from garden earth; S.<br />
basalis Zetterstedt from deciduous leaf litter; S. obsoleta Fallen from rotting vegetable<br />
matter and S. sordida Haliday from wrens' nests. Larval habitats <strong>of</strong> <strong>the</strong> remaining six<br />
<strong>British</strong> species are unrecorded and none <strong>of</strong> <strong>the</strong> larvae have been described. Some larval<br />
details <strong>of</strong> <strong>the</strong> European S. quadripunctata (L.) are very poorly illustrated by Perris<br />
( 1852) who found <strong>the</strong> larva in rotting straw.<br />
Peplomyza litura (Meigen) has been reared from a larva mining <strong>the</strong> wi<strong>the</strong>red leaf <strong>of</strong><br />
crab apple and <strong>the</strong> larval mouthparts are here illustrated from a puparium among A. H.<br />
Hamm's material in <strong>the</strong> Hope En<strong>to</strong>mological Collections at Oxford (fig. 400).<br />
Aulogastromyia anisodactyla (Loew) and <strong>the</strong> rare Cnemacantha muscaria (Fallen)<br />
are unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Six <strong>of</strong> <strong>the</strong> <strong>10</strong> <strong>British</strong> Lyciella species have been reared as follows: L. a./finis<br />
(Zetterstedt) from a rotting birch stump; L. decempunctata (Fallen) from thrush and<br />
blackbird nests and thistle (Carlina) heads; L. decipiens (Loew) from rotting leaves; L.<br />
pallidiventris (Fallen) under rotten bark, garden earth and flood refuse; L. rorida<br />
(Fallen) from rotting leaves, under bark <strong>of</strong> decayed tree trunk, earth, moss on rocks;<br />
L. subfasciata (Zetterstedt) from rotting tree leaves. L. rorida (figs 396--399) is <strong>the</strong> only<br />
<strong>British</strong> species for which <strong>the</strong> <strong>immature</strong> <strong>stages</strong> have been described (Hennig, 1952).<br />
Tricholauxania praeusta (Fallen) has been reared from rotting tree leaves and() cow<br />
dung.<br />
Larvae <strong>of</strong> Calliopum aeneum (Fallen) have been found in Viola ovaries; leaves, stems,<br />
bases and root-collars <strong>of</strong> clover (Trifolium); rotting tree leaves and crab apple leaflitter.<br />
C. simillimum (Collin) has been reared from a robin's nest. Nothing is known <strong>of</strong> <strong>the</strong> two<br />
remaining <strong>British</strong> Calliopum species and none has been described in <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong>.<br />
Lauxania cylindricornis (F.) has been reared from witches' broom on black spruce<br />
(Picea mariana (Miller)) in Canada and Teskey (1976) gives a brief unillustrated<br />
description <strong>of</strong> <strong>the</strong> larva.<br />
Of <strong>the</strong> 5 <strong>British</strong> species <strong>of</strong> Homoneura only H. notata (Fallen) has been reared from<br />
larvae found in rotting tree leaves. Several Nearctic species have been reared from<br />
similar sources (Acer, Prunus, Alnus, etc.) and <strong>the</strong> larva <strong>of</strong> H. americana (Wiedemann)<br />
(Nearctic) is illustrated here (figs 403-404).<br />
Coelopidae<br />
(Figs: larvae 406-412, puparium 1181)<br />
The small family Coelopidae (seaweed flies) contains 20 species in about <strong>10</strong> genera<br />
and is represented in all regions except <strong>the</strong> Oriental. The larvae develop in decaying<br />
93
sea weeds on seashores. Although adults may sometimes occur inland in large swarms,<br />
attracted <strong>to</strong> particular chemicals (e.g. trichlorethylene), it is doubtful if <strong>the</strong>y ever breed<br />
away from <strong>the</strong>ir special seashore niche (Oidroyd, 1954). Only 3 species in 2 genera<br />
occur in Britain.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Malacomyia sciomyzina (Haliday) are unknown but this<br />
species undoubtedly breeds in rotting seaweed where <strong>the</strong> adults occur sparingly in<br />
mostly sou<strong>the</strong>rn coastal localities.<br />
Egglishaw (1960a) and Burnet (1961) have studied <strong>the</strong> biology <strong>of</strong> <strong>the</strong> two <strong>British</strong><br />
species Coelopa frigida (F.) (figs 406-409) and C. pilipes Haliday (figs 41 0--412). Larvae<br />
<strong>of</strong> both species occur in wrack banks ra<strong>the</strong>r than <strong>the</strong> wrack string, though rarely<br />
<strong>to</strong>ge<strong>the</strong>r and <strong>the</strong>n only a few <strong>of</strong> C. pilipes will be present. C.frigida larvae, however, are<br />
never found among a large number <strong>of</strong> C. pilipes.<br />
In spite <strong>of</strong> several detailed studies on wrack fauna (e.g. Backlund, 1945b,<br />
Egglishaw, 196Q-1961) <strong>the</strong> establishment <strong>of</strong> <strong>the</strong> precise ecological requirements <strong>of</strong> each<br />
species <strong>of</strong> <strong>the</strong> several families involved (Coelopidae, Helcomyzidae, Sepsidae<br />
(Orygma)) still needs refinement and <strong>the</strong> subject is an excellent one for <strong>the</strong> purposes <strong>of</strong><br />
research, teaching and demonstration.<br />
Work on Coelopa genetics, started at Newcastle in <strong>the</strong> late 1950s by Dr Ursula Phi lip,<br />
was continued by Dr Barrie Burnet who is now at Sheffield. Genetic and o<strong>the</strong>r studies<br />
are also in progress at Nottingham and yielding information on <strong>the</strong> biology <strong>of</strong> <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> <strong>of</strong>Ceolopidae (e.g. Butlin & Day, 1984).<br />
Heleomyzidae<br />
(Figs: larvae 413-446, puparia 1182-1183)<br />
This is a cosmopolitan family <strong>of</strong> over 500 species in some 65 genera <strong>of</strong> which 56<br />
species in 16 genera occur in Britain. The classification and nomenclature <strong>of</strong> <strong>the</strong> family<br />
has been considerably revised since <strong>the</strong> <strong>British</strong> Check List appeared. Gorodkov (in<br />
Soos & Papp, 1984) should now be consulted. Following Gorodkov <strong>the</strong> genera are<br />
grouped here under 3 subfamilies but since so few <strong>immature</strong> <strong>stages</strong> are known his tribal<br />
classification is omitted.<br />
The fungus hosts <strong>of</strong> <strong>the</strong> family are fully listed by Chandler (in Stubbs & Chandler,<br />
1978). Hackman (1963, 1967) records adult Heleomyzidae found in <strong>the</strong> burrows and<br />
nests <strong>of</strong> small mammals and Hicks (1959- 1971) lists those found in birds' nests. <strong>An</strong><br />
interesting review <strong>of</strong> <strong>the</strong> larval habits <strong>of</strong>Nearctic Heleomyzidae is given by Garnett &<br />
Foote ( 1967). A key <strong>to</strong> some species (in Russian) is provided by Lobanov (1970c).<br />
Heleomyzinae. The larvae <strong>of</strong> this subfamily are presumably saprophagous and have<br />
been recorded from carrion, faeces, mammals' burrows and birds' nests; <strong>the</strong>y also<br />
occur in caves.<br />
Heleomyza serrata (L.) (figs 413--418) has been reared from <strong>the</strong> fungus Fistulina but is<br />
usually found in o<strong>the</strong>r media, e.g. birds' nests, caves and soil.<br />
Neoleria ruficauda (Zetterstedt) visits fungi and carrion and N. inscripta (Meigen)<br />
(figs 419--423) although an uncommon fly was reared in numbers from a dead fox<br />
(Smith, 1975).<br />
Scoliocentra villosa (Meigen) (figs 424--428) has been reared from dead hens at <strong>the</strong><br />
entrance <strong>to</strong> a fox-hole (Skidmore, 1967) and Seguy (1934) records <strong>the</strong> fly from bat<br />
roosts and rabbit holes. The adult has been found in every month <strong>of</strong> <strong>the</strong> year, frequently<br />
in caves, which has earned it <strong>the</strong> common name <strong>of</strong> cave-fly.<br />
Oeco<strong>the</strong>afenestralis (Fallen) (figs 429--434) is ano<strong>the</strong>r carrion feeding species found<br />
in mammal burrows and caves.<br />
Suillinae. Larvae <strong>of</strong> this subfamily mostly develop in fungi but on <strong>the</strong> Continent<br />
Suillia lurida Meigen is a pest <strong>of</strong> onion and garlic and is figured here (figs 435--436) as it<br />
94
may be imported. S. humilis (Meigen), S. pa/lida (Fallen) and S. ustulata (Meigen) have<br />
been reared from truffles (Tuber). S. bicolor (Zetterstedt), S. fuscicornis (Zetterstedt)<br />
and S. variegata (Loew) are polyphagous on many agarics, boleti, etc. O<strong>the</strong>r species<br />
appear <strong>to</strong> be more restricted in <strong>the</strong> range <strong>of</strong> <strong>the</strong>ir fungus hosts but some have been<br />
reared from o<strong>the</strong>r pabula, e.g. S. ustulata from dead elder stem and S. variegata from<br />
roots <strong>of</strong> Aster.<br />
Heteromyzinae. Heteromyza oculata Fallen has been reared from an unidentified<br />
lignicolous fungus.<br />
Tephrochlamys .flavipes (Zetterstedt) is recorded from several agarics, s<strong>of</strong>t<br />
polypores, Tuber and bird and mammal nests; T. tarsalis (Zetterstedt) (figs 437-441)<br />
from several agarics and birds' nests; T. rufiventris (Meigen) (figs 442-446), one <strong>of</strong> <strong>the</strong><br />
commonest species <strong>of</strong> <strong>the</strong> family, from Hypoxylon fragiforme and a wide range <strong>of</strong><br />
decaying animal or vegetable matter.<br />
Trixoscelidae<br />
This small family was formerly included in <strong>the</strong> Heleomyzidae and is still regarded as<br />
such by some workers. It contains some 70 species mostly found in <strong>the</strong> Holarctic and<br />
Afrotropical regions. Nothing is known <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> but Trixoscelis<br />
canescens (Loew) was reared from a blackbird's nest by <strong>the</strong> late E. B. Basden and Dr<br />
A. G. Irwin has reared a species from soil derived from feral pigeon guano. Two <strong>of</strong> <strong>the</strong><br />
four <strong>British</strong> species (T. obscurella (Fallen) and T. marginella (Fallen)) appear <strong>to</strong> have a<br />
coastal distribution (seep. 9).<br />
Chyromyidae<br />
<strong>An</strong>o<strong>the</strong>r small family (about 40 species) which is represented in most regions and in<br />
Britain by 6 species in 3 genera. The <strong>immature</strong> <strong>stages</strong> are undescribed but <strong>the</strong> larvae are<br />
said <strong>to</strong> develop in guano near <strong>the</strong> nests or roosting places <strong>of</strong> birds, in mammal burrows<br />
and under <strong>the</strong> bark <strong>of</strong> trees. Dr A. G. Irwin has reared a Chyromya from damp<br />
droppings <strong>of</strong> Dauben<strong>to</strong>n's bat.<br />
Ardo ( 1957) regards Chyromya minima (Becker) as restricted <strong>to</strong> sand-dunes and<br />
Se guy ( 1934) regards C. .flava (L.), C. oppidana (Scopoli) and Gymnochyromyia inermis<br />
Collin as cave-dwellers (troglobionts) and birds' nests dwellers (nidicoles). Hicks<br />
( 1959) records C. flava and G. inermis from birds' nests ('owl'; and wren, hedgesparrow,<br />
blackbird, greenfinch, etc., respectively). Adults <strong>of</strong> Aphaniosoma socium<br />
Collin and A. propinquans Collin are recorded from sea-shore flowers (Convolvulus and<br />
Matricaria).<br />
Sepsidae<br />
(Figs: larvae 447--474, eggs <strong>10</strong>31-<strong>10</strong>33, puparia 1184-1185)<br />
This family contains some 240 species in 21 genera and is well represented in all <strong>the</strong><br />
zoogeographical regions. Their wide distribution is undoubtedly due <strong>to</strong> <strong>the</strong>ir close<br />
association with animal faeces in which <strong>the</strong>y mostly breed. Twenty-seven species in 6<br />
genera are known from <strong>the</strong> <strong>British</strong> Isles.<br />
The larvae <strong>of</strong> several species have been described by Hennig (1949), Schumann<br />
(1962), Brindle (1965b), Mangan (1977) and Ozerov (1986). Pant's (1979) Handbook <strong>to</strong><br />
adults includes a substantial section on <strong>the</strong> life his<strong>to</strong>ries and <strong>immature</strong> <strong>stages</strong>.<br />
Orygma luctuosum Meigen (figs 447--450) breeds on <strong>the</strong> sea-shore in <strong>the</strong> smaller drier<br />
beds <strong>of</strong> decomposing wrack and was until recently included in <strong>the</strong> Coelopidae.<br />
95
Saltella sphondylii (Schrank) (figs 451-453) breeds in cow dung.<br />
Little is known <strong>of</strong> <strong>the</strong> biology and <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> 11 <strong>British</strong> species <strong>of</strong><br />
Themira, but probably most develop in mud alongside ponds and streams, especially<br />
if this is enriched with dung or sewage effluent. T. putris (L.) (figs 454-456) and<br />
T. nigricornis (Meigen) have been recorded from human excrement and <strong>the</strong> former<br />
breeds in <strong>the</strong> sludge and seepage from sewage works and can produce vast numbers<br />
<strong>of</strong> adults which may become a nuisance by invading nearby houses. Larvae <strong>of</strong><br />
T. annu/ipes Meigen occur in soil especially if enriched by cattle droppings, sewage<br />
overflows, etc. T. minor Haliday occurs in cow dung and trampled dung and soil from<br />
cattle feeding pens. T. /eachi (Meigen) probably breeds in cow dung upon which <strong>the</strong><br />
females may frequently be seen and it has been reared from manured soil, human<br />
excrement and grass cuttings. T. nigricornis has also been reared from garden soil and<br />
dung (human and chicken) and T. putris has been reared from birds' nests. Adults <strong>of</strong><br />
T. biloba <strong>An</strong>derson, recently added <strong>to</strong> <strong>the</strong> <strong>British</strong> List (Pont, 1986) were swept <strong>of</strong>f<br />
guano on a coot's nest. T. lucida Staegar and T. superba Haliday seem also <strong>to</strong> be<br />
associated with waterfowl droppings though <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are unknown.<br />
Abroad several species have been reared from <strong>the</strong> dung <strong>of</strong> cows, pigs, humans and<br />
small mammals.<br />
The rare Meroplius minutus Wiedemann ( =stercorarius Robineau-Desvoidy) (figs<br />
457-460) has been reared from human faeces, cow, pig and (accumulations <strong>of</strong>) rabbit<br />
dung.<br />
Nemopoda nitidu/a (Fallen) (figs 461-465) has been reared from human excrement,<br />
dead snails, carrion and (mainly rotting) fungi.<br />
All Sepsis species (figs 466-474) breed in mammal dung, preferably cow dung, but<br />
also including horse dung (S. fu/gens Meigen, S. violacea Meigen), chicken dung<br />
(S. vio/acea, S.fulgens), pig dung (S.fu/gens, S. punctum (F.), S. vio/acea), human faeces<br />
(S. punctum), sewage and pig swill (S.fu/gens), sheep dung (S. cynipsea (L.), in upland<br />
areas), small mammal droppings (S. punctum, S. neocynipsea Melander & Spuler, <strong>the</strong><br />
latter also allegedly in small mammal carrion).<br />
The eggs <strong>of</strong>Sepsidae (figs <strong>10</strong>31-<strong>10</strong>33) have long respira<strong>to</strong>ry horns (Orygma has 2)<br />
<strong>of</strong>ten much longer than <strong>the</strong> egg itself. All are plastron bearing (except Saltella) and are<br />
adapted for atmospheric respiration and for extracting dissolved oxygen from ambient<br />
water when <strong>the</strong> egg is submerged (Hin<strong>to</strong>n, 1960, 1961, 1981 ).<br />
Sciomyzidae<br />
(Figs: larvae 475-518, eggs <strong>10</strong>34--<strong>10</strong>36, puparia 1186-1189, cover)<br />
Over 470 species <strong>of</strong> Sciomyzidae (snail killing flies) have been described throughout<br />
<strong>the</strong> world. C. 0. Berg & Knutson ( 1978) provide a useful summary <strong>of</strong> <strong>the</strong> state <strong>of</strong><br />
knowledge <strong>of</strong> <strong>the</strong> biology and systematics <strong>of</strong> <strong>the</strong> family which has increased rapidly<br />
over <strong>the</strong> past 30 years.<br />
The discovery that <strong>the</strong>ir larvae feed on living aquatic and terrestrial molluscs gave<br />
<strong>the</strong>m a medical and veterinary significance in <strong>the</strong> possible control <strong>of</strong> schis<strong>to</strong>somiasis (in<br />
Africa, South America and <strong>the</strong> Far East) and o<strong>the</strong>r snail-borne diseases. The natural<br />
food <strong>of</strong> over 200 species is now known and <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> many species have<br />
been described in detail.<br />
The larval morphology reflects <strong>the</strong>ir mode <strong>of</strong> life. Free-living larvae have wart-like<br />
tubercles on <strong>the</strong> body and elongate lobes on <strong>the</strong> anal segments. Aquatic forms have<br />
conspicuous branched float hairs on <strong>the</strong> posterior spiracular plate. Larvae living as<br />
parasi<strong>to</strong>ids have a mostly transparent integument without conspicuous tubercles but<br />
with a transverse patch <strong>of</strong> spinules on each segment <strong>of</strong> <strong>the</strong> body.<br />
Sixty-five species in 23 genera representing 3 subfamilies occur in Britain. Rozkosny<br />
(1967) provides keys <strong>to</strong> larvae and puparia <strong>of</strong> some genera and species.<br />
96
Salticellinae. Salt ice/la fascia la (Meigen) (figs 475-479) was <strong>the</strong> first species <strong>of</strong> <strong>the</strong><br />
family <strong>to</strong> be associated with molluscs when Perris (1850) reared adults from puparia<br />
found in <strong>the</strong> shells <strong>of</strong> Theba pisana (Miiller). Knutson, Stephenson & Berg (1970)<br />
described <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and recorded <strong>the</strong>m from several species <strong>of</strong> snails found<br />
on dunes or in dry habitats (e.g. Theba pisana, He/ice/la candicans (Pfieffer), H. virgata<br />
(de Costa) and H. hortensis (Miiller), all Helicellidae) on which <strong>the</strong>y may feed as<br />
solitary, internal parasi<strong>to</strong>ids or <strong>the</strong>y may be saprophagous on dead tissue.<br />
Phaeomyiinae. Nothing is published on <strong>the</strong> biology or <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> two<br />
<strong>British</strong> species <strong>of</strong> Pe/idnoptera, but Dr Peter Baily has recently reared a species from<br />
millipedes in Portugal.<br />
Sciomyzinae<br />
Sciomyzini. Co/obaea bifascie/la (Fallen) has a highly specialized parasi<strong>to</strong>id larva<br />
which feeds as a solitary individual and each larva consumes only one snail. One or two<br />
eggs are laid across <strong>the</strong> sutures <strong>of</strong> <strong>the</strong> shell <strong>of</strong> exposed Lymnaea palustris Miiller or<br />
L. trunculata Miiller. The larva enters <strong>the</strong> shell and feeds on <strong>the</strong> mucus, extra pallial<br />
fluid, or <strong>the</strong> less vital tissues. Later <strong>the</strong> vital tissues are consumed, <strong>the</strong> snail dies and <strong>the</strong><br />
larva consumes <strong>the</strong> decaying tissues; <strong>the</strong>n, after about 25 days, it pupates within <strong>the</strong><br />
shell. The adult emerges some two or three weeks later. Although more than one egg<br />
may be laid only one larva feeds and it is thought that it kills any o<strong>the</strong>r larvae as <strong>the</strong>y<br />
enter <strong>the</strong> shell.<br />
C. puncta/a (Lundbeck) has been recorded from Planorbis a/bus Miiller, Lymnaea<br />
peregra (Miiller) and young P. corneus (L.). C. pec<strong>to</strong>ralis (Zetterstedt) and C. distincta<br />
(Meigen) (fig. 480) have been reared from <strong>An</strong>isus vortex (L.) and A. spirorbis (L.)<br />
respectively.<br />
Of <strong>the</strong> 16 species <strong>of</strong> Pherbellia occurring in Britain <strong>the</strong> life his<strong>to</strong>ries and <strong>immature</strong><br />
<strong>stages</strong> <strong>of</strong>9 species have been described (Bratt et al. 1969). Eggs are laid in mosses, etc.<br />
and <strong>the</strong> larvae seek out and may eat up <strong>to</strong> 4 snails during <strong>the</strong>ir development.<br />
Most <strong>of</strong> <strong>the</strong> common species <strong>of</strong> Pherbellia feed on hygrophilous and exposed aquatic<br />
snails, whereas <strong>the</strong> uncommon species eat terrestrial snails.<br />
Some snails eaten in natural conditions are indicated as follows:- Terrestrial species:<br />
Pherbe/lia dubia (Fallen) (figs 482-483) (Cochlicopa, Discus, Hygromia, Oxychilus,<br />
Retine/la); P. knutsoni Verbeke (Coch/ice/la, He/ice/la); P. scute/laris (von Roser)<br />
( C/ausi/ia). Aquatic and hygrophilous species: Pherbe/lia dorsal a (Zetterstedt)<br />
(Planorbis); P. schoenherri (Fallen) (fig. 481) (Succinea); P. scute/laris and P. ventralis<br />
(Fallen) (figs 484-485) (Lymnaea). Full lists including snails eaten or rejected in labora<strong>to</strong>ry<br />
conditions are given by Bratt et al. (1969) and 0. Beaver (1972) gives fur<strong>the</strong>r<br />
information on some <strong>British</strong> species.<br />
The larvae <strong>of</strong> Pteromicra species are polyphagous preda<strong>to</strong>rs <strong>of</strong> small aquatic and<br />
hygrophilous, non-operculate snails and may kill and consume up <strong>to</strong> 3 snails during<br />
<strong>the</strong>ir development. The common P. angustipennis (Staeger) (figs 486-489) feeds on<br />
Planorbis con<strong>to</strong>rtus (L.), young P. planorbis (L.), P. vortex (L.), Hygromia hispida (L.),<br />
Succinea sp., Segmentina nitida (Miiller), and Lymnaea truncatu/a (Miiller). Pteromicra<br />
glabricu/a (Fallen) feeds on Planorbis vortex, P. con<strong>to</strong>rtus, young individuals <strong>of</strong><br />
Physa fontina/is (L.), Planorbis planorbis, Lymnaea palustris, L. peregra (Miiller), L.<br />
truncatula, and <strong>the</strong> terrestrial snails Discus rotundatus (Miiller) and Succinea sp.<br />
Pteromicra /eucopeza (Meigen) larvae have been found in Planorbis con<strong>to</strong>rtus, and<br />
Pteromicra pec<strong>to</strong>rosa (Hendel) feeds on Segmentina nitida and Planorbis vortex . The<br />
<strong>immature</strong> <strong>stages</strong> <strong>of</strong> Pteromicra are fully described by Rozkosny & Knutson (1970).<br />
All known larvae <strong>of</strong> Sciomyza feed on terrestrial snails <strong>of</strong> <strong>the</strong> family Succineidae and<br />
<strong>the</strong> <strong>immature</strong> <strong>stages</strong> are described by Foote (1959). The two <strong>British</strong> species are known<br />
<strong>to</strong> attack living snails <strong>of</strong> <strong>the</strong> genus Oxyloma (in Alaska). Sciomyza simplex Fallen<br />
97
(figs 490-492) is probably a preda<strong>to</strong>r (several attacking more than one snail) ra<strong>the</strong>r<br />
than parasi<strong>to</strong>id (one snail per larva) but <strong>the</strong> feeding status <strong>of</strong> S. dryomyzina Zetterstedt<br />
is not yet clear.<br />
The larva <strong>of</strong> Tetanura pallidiventris Fallen (figs 493-494) feeds on terrestrial snails <strong>of</strong><br />
<strong>the</strong> genera Cochlicopa, Discus and Retinella and may have some use as a biological<br />
control agent as one <strong>of</strong> <strong>the</strong> host snails, Cochlicopa lubrica (Miiller), is an intermediate<br />
host <strong>of</strong> <strong>the</strong> sheep lancet fluke, Dicrocoelium dendriticum (Rudolphi) (Trema<strong>to</strong>da).<br />
Knutson (1970a) describes <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and biology. Eggs are laid directly on<strong>to</strong><br />
<strong>the</strong> retracted s<strong>of</strong>t parts <strong>of</strong> <strong>the</strong> host snail. The puparia vary in shape according <strong>to</strong><br />
whe<strong>the</strong>r <strong>the</strong>y were formed inside or outside <strong>of</strong> <strong>the</strong> host's shell; and those formed inside<br />
vary fur<strong>the</strong>r with <strong>the</strong> shape <strong>of</strong> <strong>the</strong> host shell. A puparium formed in <strong>the</strong> elongate shell <strong>of</strong><br />
Cochlicopa lubrica is illustrated (fig. 1188).<br />
Tetanocerini. Larvae <strong>of</strong> <strong>An</strong>tichaeta feed exclusively on <strong>the</strong> eggs <strong>of</strong> snails and appear<br />
<strong>to</strong> be <strong>the</strong> only insects <strong>to</strong> do so (though <strong>the</strong> phorids Megaselia aequalis and M . ciliata<br />
feed on slug eggs and that large and enterprising genus may well possess an as yet<br />
undetected snail egg preda<strong>to</strong>r). The eggs (fig. <strong>10</strong>36) are enclosed in a gelatinous substance<br />
and are deposited on egg capsules or egg masses <strong>of</strong> <strong>the</strong> snails. A. anal is (Meigen)<br />
larvae feed on <strong>the</strong> eggs <strong>of</strong> Lymnaea truncatula (and in <strong>the</strong> labora<strong>to</strong>ry on L. stagna/is (L.)<br />
and Succinea sp.) and A. brevipennis (Zetterstedt) (figs 495-498) feeds on egg masses <strong>of</strong><br />
Succinea sp. (and in <strong>the</strong> labora<strong>to</strong>ry in egg capsules <strong>of</strong> Lymnaea truncatula). Knutson<br />
( 1966) describes <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and detailed biology <strong>of</strong> <strong>the</strong> genus.<br />
Coremacera tristis (Harris) ( =marginata F.) (figs 499-503) has been reared on a<br />
number <strong>of</strong> terrestrial snails in <strong>the</strong> labora<strong>to</strong>ry (Knutson, 1973).<br />
Dictya umbrarum (L.) was reared by Lundbeck (1923) from floating puparia and is<br />
presumed <strong>to</strong> attack aquatic snails as do <strong>the</strong> larvae <strong>of</strong> some Nearctic species described by<br />
Valley & Berg (1977).<br />
Larvae <strong>of</strong> <strong>the</strong> genus Elgiva (figs 504--506) attack aquatic pulmonate snails <strong>of</strong> <strong>the</strong><br />
families Lymnaeidae, Physidae and Planorbidae. The larvae are air-brea<strong>the</strong>rs and float<br />
just beneath <strong>the</strong> surface film <strong>of</strong> quiet waters and <strong>the</strong>y forage in and on Lemna mats.<br />
They quickly kill snails and commence feeding and when replete move on <strong>to</strong> o<strong>the</strong>r<br />
snails; each larva may killl2- 23 snails during its development. The puparia (fig. 1187,<br />
cover) are well adapted for flotation and have <strong>the</strong> posterior spiracles uplifted. Knutson<br />
& Berg (1964) describe <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and biology <strong>of</strong> <strong>the</strong> genus.<br />
The larvae <strong>of</strong> Hydromya dorsalis (F.) (figs 507-509) are found in shallow, flowing<br />
water where <strong>the</strong>y kill and feed upon aquatic pulmonate snails, especially Lymnaea spp.<br />
(Knutson & Berg, 1963).<br />
Knutsonia species (except K. lineata) are also preda<strong>to</strong>rs <strong>of</strong> aquatic pulmonate snails<br />
and a larva may consume as many as 40 snails during <strong>the</strong> 21-77 days <strong>of</strong> its development.<br />
The larvae <strong>of</strong> both <strong>British</strong> species, K. albiseta (Scopoli) and K. lineata (Fallen) (figs<br />
5<strong>10</strong>--511) are described by Knutson & Berg (1967) but <strong>the</strong>y were unable <strong>to</strong> rear K.<br />
lineata on gastropod molluscs and it has since been established (Foote & Knutson,<br />
1970) that this species is exceptional in feeding only on bivalve clams (Pisidium}, a habit<br />
also shared by <strong>the</strong> genus Renocera (see below). During <strong>the</strong> 37- 53 days <strong>of</strong> larval life<br />
K. lineata killed and ate 21- 31 individual clams (1 ·5-4·0 mm in greatest diameter).<br />
The larvae <strong>of</strong> Limnia, Pherbina and Psacadina are predaceous on hygrophilous and<br />
exposed aquatic snails at <strong>the</strong> shoreline or on <strong>the</strong> water surface. Knutson, Rozkosny &<br />
Berg ( 1975) describe <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Pherbina and Psacadina. Pherbina cory leti<br />
(Scopoli}, one <strong>of</strong> <strong>the</strong> commonest European sciomyzids, is illustrated here (figs 512-<br />
514).<br />
The larva <strong>of</strong> Renocera striata (Meigen) is an aquatic preda<strong>to</strong>r <strong>of</strong> sphaeriid clams. The<br />
o<strong>the</strong>r two <strong>British</strong> species may also be presumed <strong>to</strong> be so as Foote ( 1976) has shown that<br />
<strong>the</strong> habit is shared by American species. Renocera and Knutsonia lineata (discussed<br />
above) are <strong>the</strong> only insects known <strong>to</strong> kill and feed on bivalve molluscs. The only o<strong>the</strong>r<br />
98
insect found associated with clams is <strong>the</strong> larva <strong>of</strong> <strong>the</strong> chironomid Glyp<strong>to</strong>tendipes paripes<br />
(Edwards) found between <strong>the</strong> shell and mantle <strong>of</strong> <strong>An</strong>odonta cygnea L. (Beedham, 1966).<br />
Sepedon larvae are preda<strong>to</strong>rs <strong>of</strong> aquatic snails and <strong>the</strong> two <strong>British</strong> species are<br />
included in Neff & Berg ( 1966).<br />
<strong>British</strong> representatives <strong>of</strong> <strong>the</strong> genus Tetanocera include one slug-killing species, T.<br />
elata (F.) (fig. 515), and two North American species also share this habit (Knutson,<br />
Stephenson & Berg, 1965; Rozkosny, 1965; Stephenson & Knutson, 1966). O<strong>the</strong>r<br />
members <strong>of</strong> <strong>the</strong> genus range over <strong>the</strong> usually snail-killing habits <strong>of</strong> <strong>the</strong> family as<br />
follows: T. ferruginea Fallen (figs 516-518), T. hyalipennis von Roser, T. robusta<br />
Loew (aquatic snails); T. arrogans (Meigen), T. silvatica Meigen, T. unicolor Loew<br />
(hygrophilous and exposed aquatic snails); T. phyllophora Melander (terrestrial snails).<br />
The biology <strong>of</strong> T. punctifrons Rondani requires description and fur<strong>the</strong>r elucidation<br />
(0. Beaver, 1972).<br />
The biology and <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Trype<strong>to</strong>ptera punctulata (Scopoli) remain<br />
unknown.<br />
Nothing is known <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> two <strong>British</strong> species <strong>of</strong> Diche<strong>to</strong>phora<br />
or <strong>of</strong> Ectinocera borealis (Zetterstedt).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Euthycerafumigata (Scopoli) are unknown but <strong>the</strong> larva may<br />
attack slugs as <strong>the</strong> European species E. chaerophyli (F.) appears <strong>to</strong> do (Knutson, 1970b)<br />
(see also under Tetanocera above).<br />
Much is known <strong>of</strong> <strong>the</strong> biology and life-his<strong>to</strong>ries <strong>of</strong> this fascinating family but much<br />
detail still needs <strong>to</strong> be added, particularly on which species <strong>of</strong> snail are selected as food<br />
in <strong>the</strong> field ra<strong>the</strong>r than in <strong>the</strong> labora<strong>to</strong>ry.<br />
Techniques for labora<strong>to</strong>ry rearing are described by Willomitzer (1970) and<br />
Willomitzer & Rozkosny ( 1977). The distribution <strong>of</strong> <strong>the</strong> family in Britain is given by<br />
Stephenson & Knutson (1970).<br />
Sphaeroceridae<br />
(Figs: larvae 519-530, eggs <strong>10</strong>37-<strong>10</strong>42, puparia 1190-1193)<br />
About <strong>10</strong>00 species <strong>of</strong> Sphaeroceridae have been described from all regions <strong>of</strong> <strong>the</strong><br />
world. Many anthropophilic species have a wide distribution having been introduced<br />
by man in<strong>to</strong> continents in which <strong>the</strong>y are not endemic. Their propensity for breeding in<br />
dung no doubt enabled <strong>the</strong> flies <strong>to</strong> follow caravan routes overland and shipping routes<br />
in <strong>the</strong> holds <strong>of</strong> cargo ships carrying cattle and o<strong>the</strong>r domestic animals. The larvae are<br />
generally saprophagous and in addition <strong>to</strong> dung will develop in mud, forest litter, fungi,<br />
decaying vegetable refuse, seaweed, in caves, in <strong>the</strong> runs <strong>of</strong> mammals, in <strong>the</strong> nests <strong>of</strong><br />
birds, mammals and insects, and in carrion.<br />
Some 114 species in 33 genera and 3 subfamilies occur in Britain. The <strong>immature</strong><br />
<strong>stages</strong> are not well known and <strong>the</strong>re are more descriptions <strong>of</strong> <strong>the</strong> distinctive puparia<br />
(figs 1190-1193), (Goddard, 1938; Okely, 1974) than any o<strong>the</strong>r stage. Puparia are<br />
frequently found during archaeological excavations (see section on fossils). Some<br />
larvae have been described by Schumann (1962), and Pitkin (1987) reviews and<br />
reproduces illustrations <strong>of</strong> most <strong>of</strong> <strong>the</strong> described <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> family in his<br />
Handbook <strong>to</strong> <strong>the</strong> <strong>British</strong> species. Carrion and dung frequenting species <strong>of</strong> possible<br />
forensic significance are discussed by Smith (1986b). <strong>An</strong> American species has been<br />
involved in a case <strong>of</strong> human intestinal myiasis (Micks & McKibben, 1956).<br />
The following comments are confined <strong>to</strong> actual field rearing records since <strong>the</strong> adults<br />
are frequently found on media in which <strong>the</strong>y do not necessarily breed. Pitkin (1987)<br />
should be consulted for a review <strong>of</strong> adult habits. In <strong>the</strong> labora<strong>to</strong>ry boiled grass-cuttings<br />
have proved a successful medium on which <strong>to</strong> rear larvae when <strong>the</strong>ir natural pabulum is<br />
unknown.<br />
99
Sphaerocerinae. Sphaerocera curvipes Latreille (fig. 519) is a common species that has<br />
been reared from cow dung and human excrement. lschiolepta pus ilia (Fallen) has been<br />
reared from human excrement and dead snails. Lo<strong>to</strong>bia pallidiventris (Meigen) has been<br />
reared from cow-dung.<br />
Copromyzinae. Lo<strong>to</strong>phila ( = Copromyza) atra (Meigen) (figs 520-522) has been<br />
reared from cow-dung. Crumomyia nitida has been reared from horse-dung and fungi;<br />
C. pedestris (Meigen) has been reared from dead snails. Copromyza similis (Collin)<br />
has been reared from cow-dung, C. equina Fallen from horse and cow-dung and C.<br />
stercoraria (Meigen) from human excrement.<br />
Limosininae. The following species breed in cow dung: Coproica acutangula<br />
(Zetterstedt) (figs 523-524), C. lugubris (Haliday), C. vagans (Haliday), Elachisoma<br />
aterrima (Haliday), Halidayina spinipennis (Haliday), Chae<strong>to</strong>podella scutellaris<br />
(Haliday) (figs 525- 526), Limosina silvatica (Meigen), Opalimosina collini (Richards),<br />
0 . denticulata (Duda), Spelobia clunipes (Meigen), and Telomerina pseudoleucoptera<br />
(Duda). Of <strong>the</strong>se, Coproica vagans and Limosina silvatica (fig. 527) have also been<br />
reared from human excrement and horse dung respectively.<br />
Lep<strong>to</strong>cera caenosa (Rondani) breeds in human excrement and is particularly<br />
associated with water closets where it can occur in large numbers causing a considerable<br />
nuisance and health hazard (Fredeen & Taylor, 1964).1t has also been reared from<br />
wasps' nests.<br />
Lep<strong>to</strong>cera fontinalis (Fallen), Spelobia palmata Richards, S. nana (Rondani) and<br />
S. luteilabris (Rondani) have been reared from dead snails. Spelobia parapusio (Dahl)<br />
(fig. 528) has been reared from fungi. Apteromyia claviventris (Strobl) and Minilimosina<br />
fungicola (Haliday) have occurred in a nest <strong>of</strong> <strong>the</strong> ant Lasius fuliginosus (L.). The<br />
former species has been found in roots <strong>of</strong> rhubarb and, along with Pullimosina<br />
heteroneura (Haliday), has also been reared from damaged narcissus bulbs. Kimosina<br />
empirica (But<strong>to</strong>n) has been reared from a dead seal, rabbit corpses, a dead human and<br />
in sewage disposal tanks.<br />
Terrilimosina racovitzai (Bezzi) is a cave-dwelling species that may also occur<br />
indoors in cellars and has bred in a blocked drain (see introduc<strong>to</strong>ry section on caves).<br />
Thoracochaeta zosterae (Haliday) (figs 529- 530) breeds in seaweed as do o<strong>the</strong>r<br />
members <strong>of</strong> <strong>the</strong> genus; Egglishaw (1961) describes <strong>the</strong> <strong>immature</strong> <strong>stages</strong> in detail.<br />
Pallopteridae<br />
(Figs: larvae 531-538, puparium 1194)<br />
This is a small poorly studied family <strong>of</strong> only 44 species contained in 8 genera <strong>of</strong> which<br />
11 species in 2 genera occur in Britain.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Palloptera usta (Meigen) and P. ustulata Fallen (figs 531-<br />
535) were described by Morge ( 1956) from larvae found under <strong>the</strong> bark <strong>of</strong> dead coniferous<br />
and deciduous trees in Europe where <strong>the</strong>y were preying upon <strong>the</strong> larvae <strong>of</strong><br />
Cerambycidae and Scolytidae (Coleoptera). Morge ( 1967) has found that <strong>the</strong> larvae<br />
may exert a considerable control <strong>of</strong> mass outbreaks <strong>of</strong> bark beetles, especially in<br />
mountain forests. Larvae <strong>of</strong> a North American species have also been reported from<br />
<strong>the</strong> galleries <strong>of</strong> scolytid beetles and those <strong>of</strong> a Canadian species prey upon cecidomyiid<br />
larvae in Douglas fir cones (Teskey 1976).<br />
Smith (1957a) found larvae <strong>of</strong> P. ustulata under poplar bark and puparia <strong>of</strong> P.<br />
quinquemaculata Macquart (figs 536-538) in soil under fool's parsley (Aethusa cynapium<br />
L.) and dandelion (Taraxacum).<br />
Nye ( 1958) found third ins tar larvae <strong>of</strong> Palloptera quinquemaculata overwintering in<br />
basal tillers <strong>of</strong> Arrhena<strong>the</strong>rum elatius (L.) (Gramineae) and Stubbs (1969) records<br />
rearing and adult associations with Compositae and Umbelliferae.<br />
<strong>10</strong>0
Nothing is known <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ry <strong>of</strong> <strong>the</strong> rare Eurygnathomyia bicolor (Zetterstedt},<br />
so far recorded only from Yorkshire.<br />
Lonchaeidae<br />
(Figs: larvae 539-547, cover, egg <strong>10</strong>43, puparia 1195- 1196)<br />
Some 300 species in I 0 genera representing <strong>the</strong> two subfamilies <strong>of</strong> this family have<br />
been described from all forested regions <strong>of</strong> <strong>the</strong> world except New Zealand. Of <strong>the</strong>se, 29<br />
species in 3 genera in two subfamilies occur in Britain. They are not well known in <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> but Smith (1957b}, Morge (1963) and Mamaev et al. (1977) illustrate<br />
some <strong>of</strong> <strong>the</strong> species occurring in Britain.<br />
Dasiopinae. The larvae <strong>of</strong> some Dasiops species live in galls formed on grasses and<br />
o<strong>the</strong>rs are found under <strong>the</strong> bark <strong>of</strong> dead or dying trees, but no <strong>British</strong> species appear <strong>to</strong><br />
have been reared.<br />
Lonchaeinae. Earomyia schis<strong>to</strong>pyga Collin feeds on <strong>the</strong> seeds <strong>of</strong> conifers and some<br />
continental species are important pests. O<strong>the</strong>r continental species <strong>of</strong> Earomyia live in<br />
male cones <strong>of</strong> spruce or in floscules <strong>of</strong> Cirsium. The lonchaeid larva recorded by<br />
Red fern ( 1968) from Cirsium vulgare heads may belong <strong>to</strong> this genus, though it was<br />
apparently predaceous.<br />
Most Lonchaea species (figs 539- 547, cover) develop under <strong>the</strong> bark <strong>of</strong> dead and<br />
dying trees or in decomposed wood: e.g. L. britteni Collin, beech; L. fugax Becker,<br />
poplar; L. limatula Collin, beech; L. palposa Zetterstedt (figs 540-542}, birch,<br />
hawthorn, poplar; L. sylvatica Beling, poplar. Some species appear <strong>to</strong> be more or less<br />
specifically associated with particular tree species, e.g. L. collini Hackman (figs<br />
543- 544) is usually found on pines and is a facultative preda<strong>to</strong>r <strong>of</strong> scolytid beetles;<br />
L. peregrina Becker occurs under bark <strong>of</strong> Salix and poplars. Lonchaea chorea (F.)<br />
( = flavidipennis Zetterstedt) is a synanthropic species which develops in various<br />
decaying organic matter and it has been recorded from beetroot, but it is possible that<br />
only diseased plants (e.g. eelworm infested) are attacked and that <strong>the</strong> larvae are introduced<br />
in 'manure' (Farsky, 1879; Cameron, 1913). I have also seen L. chorea reared<br />
from 'stratified Rosa canina seed' (K. M. Harris).<br />
Setisquamalonchaeafumosa Egger has been reared from parsnips, turnips, and from<br />
<strong>the</strong> stems <strong>of</strong> parsley and henbane.<br />
Piophilidae<br />
(Figs: larvae 548-563, egg I 044, puparia 1197- 1199)<br />
The family Piophilidae as now unders<strong>to</strong>od (McAipine, 1977) includes <strong>the</strong> former<br />
families Neottiophilidae and Thyreophoridae and contains 67 world species in 23<br />
genera and 2 subfamilies. Thirteen species representing 4 genera and both subfamilies<br />
occur in Britain. The subfamilies are separated in <strong>the</strong> key <strong>to</strong> families (p. 71).<br />
Neottiophilinae. Only two <strong>British</strong> species <strong>of</strong> this subfamily are known, <strong>the</strong><br />
Palaearctic Neottiophilum praeustum (Meigen) and <strong>the</strong> Holarctic Actenoptera hilarella<br />
(Zetterstedt).<br />
Keilin (1924a), Tate (1954) and Zumpt (1965) describe <strong>the</strong> larva <strong>of</strong> N. praeustum (figs<br />
548- 551) which is an ec<strong>to</strong>parasitic bloodsucker on nestlings, usually <strong>of</strong> <strong>the</strong> smaller<br />
passerine birds forming more open nests e.g. greenfinch, chaffinch, linnet, blackbird,<br />
song-thrush, carrion-crow, etc. (with one record from a falconiform, <strong>the</strong> sparrowhawk).<br />
Unlike Pro<strong>to</strong>calliphora which also attacks nestlings, Neottiophilum is entirely<br />
<strong>10</strong>1
dependent on blood <strong>to</strong> complete its development and if present in sufficient numbers<br />
may kill <strong>the</strong> nestlings.<br />
The life-his<strong>to</strong>ry <strong>of</strong> Actenoptera is unknown.<br />
Piophilinae. Centrophlebomyiafurcata (F.) (figs 560-563) has not been recorded in<br />
Britain since 19<strong>10</strong> when Yerbury found adults at Porthcawl, Glamorgan. Previously he<br />
had found adults on a dead donkey, Mt Edgcumbe Park, Cornwall in April 1889. In<br />
Europe o<strong>the</strong>r 'Thyreophoridae' are associated with large skele<strong>to</strong>ns, e.g. horses,<br />
donkeys, mules and more rarely on dogs but none had been seen since 1850, in spite <strong>of</strong><br />
two World Wars, and were thought <strong>to</strong> be extinct. However, as <strong>the</strong> flies usually appeared<br />
on carcases at cold times <strong>of</strong> <strong>the</strong> year as <strong>the</strong> snows melted, it was thought <strong>the</strong> en<strong>to</strong>mologists,<br />
not normally in <strong>the</strong> field <strong>the</strong>n, may have overlooked <strong>the</strong>m. This may well be <strong>the</strong><br />
case as Freidberg (1981) has rediscovered C.furcata in Israel on <strong>the</strong> carcases <strong>of</strong> goats,<br />
cows and sheep during November <strong>to</strong> January. Eggs are laid on bones and <strong>the</strong> larvae<br />
feed and develop in <strong>the</strong> marrow. Freidberg has proposed <strong>the</strong> name 'bone-skipper' for<br />
this species since <strong>the</strong> larvae have <strong>the</strong> same skipping habit as Piophila. O<strong>the</strong>r species have<br />
since been discovered in Kashmir and Russia.<br />
The larva <strong>of</strong> Piophila casei (L.) (figs 552- 554) is <strong>the</strong> well-known 'cheese-skipper', a<br />
common s<strong>to</strong>red product pest on cheese, ham and bacon and consequently <strong>of</strong> cosmopolitan<br />
distribution. The common name refers <strong>to</strong> <strong>the</strong> habits <strong>of</strong> its skipping escape<br />
mechanism which <strong>the</strong> larva effects by arching its body and grasping <strong>the</strong> small anal<br />
papillae with its mouthparts. When <strong>the</strong> grip is released <strong>the</strong> larva is flung in<strong>to</strong> air <strong>to</strong> a<br />
height <strong>of</strong> 2 or 3 inches and may cover a horizontal distance <strong>of</strong> several inches. Several<br />
species <strong>of</strong> Piophila (including casei) feed on carrion in <strong>the</strong> larval <strong>stages</strong> and are usually<br />
late arrivals in <strong>the</strong> fauna! succession. However this may vary with <strong>the</strong> species and site <strong>of</strong><br />
<strong>the</strong> corpse and can give <strong>the</strong> larvae a forensic significance (Smith, 1986b ). The species<br />
most frequently associated with human corpses is P. foveolata Meigen ( =nigriceps<br />
Meigen) which is usually found in slaughter-houses, meat fac<strong>to</strong>ries and poultry farms.<br />
P. vulgaris Fallen (fig. 559) feeds on dead snails.<br />
There may have been some confusion <strong>of</strong> <strong>the</strong> species <strong>of</strong> Piophila associated with<br />
carrion in <strong>the</strong> early literature and it is worth identifying and recording any Piophilidae<br />
found on carrion especially if <strong>the</strong> time <strong>of</strong> demise <strong>of</strong> <strong>the</strong> corpse can be established. A key<br />
<strong>to</strong> larvae is provided by Brindle (1965c) and Smith (1986b) but <strong>the</strong> anterior spiracles<br />
(figs 554-558) afford a ready means <strong>of</strong> identifying <strong>the</strong> most common species. However,<br />
as always, it is best <strong>to</strong> rear samples through <strong>to</strong> <strong>the</strong> more readily identifiable adult stage<br />
during research projects.<br />
Opomyzidae<br />
(Figs: larvae 564-577, eggs I 045-<strong>10</strong>46, puparia 1200-120 I)<br />
A small family <strong>of</strong> over 40 species mostly found in <strong>the</strong> temperate regions <strong>of</strong> <strong>the</strong><br />
nor<strong>the</strong>rn hemisphere with a few species in East and South Africa. Fourteen species in<br />
two genera occur in Britain. Some larvae and life-his<strong>to</strong>ries are described by Thomas<br />
( 1933-1938) and Nye ( 1958). The known larvae live in <strong>the</strong> stems <strong>of</strong> Gramineae.<br />
Of <strong>the</strong> eight <strong>British</strong> species <strong>of</strong> Geomyza three are known in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Larvae <strong>of</strong> G. balachowskyi Mesnil (figs 567- 568) overwinter in <strong>the</strong> basal tillers <strong>of</strong><br />
Holcus lanatus L. and H. mollis L.; <strong>the</strong>y pupate in April and adults emerge in May and<br />
<strong>the</strong>re is probably a second generation. Geomyza tripunctata Fallen larvae (figs 564-566)<br />
overwinter in <strong>the</strong> basal tillers <strong>of</strong> Lolium multiflorum Lamarck and <strong>to</strong> a lesser extent in L.<br />
perenne L., Dactylis glomera/aL., Poa trivia/is L., P. annuaL. and wheat. They pupate<br />
in March, emerge in April and oviposit in early May. A second generation emerges in<br />
July, oviposits in August, giving rise <strong>to</strong> <strong>the</strong> overwintering larvae. Geomyza venusta<br />
(Meigen) is found in Bromus.<br />
<strong>10</strong>2
Larvae <strong>of</strong> three <strong>of</strong> <strong>the</strong> six species <strong>of</strong> Opomyza have been reared. Opomyza petrei<br />
Mesnil (figs 569- 571) overwinters in Holcus /ana/us and <strong>An</strong>thoxanthum odoratum L.;<br />
O.florum (F.) (figs 572-575) can be a serious pest <strong>of</strong> cereals, particularly winter wheat<br />
(especially in Russia). It differs from o<strong>the</strong>r members <strong>of</strong> <strong>the</strong> family in overwintering in<br />
<strong>the</strong> egg stage which hatches at <strong>the</strong> end <strong>of</strong> March and <strong>the</strong> larva bores in<strong>to</strong> wheat tillers.<br />
Pupation occurs during May and <strong>the</strong> adult emerges in early June. Oviposition occurs<br />
again in Oc<strong>to</strong>ber and early November. Eggs are laid on <strong>the</strong> soil close <strong>to</strong> <strong>the</strong> winter<br />
wheat. The wild hosts are not known. Opomyza germinationis (L.) (figs 576-577) overwinters<br />
as a third instar larva in <strong>the</strong> basal tillers <strong>of</strong> Festuca, Lolium, Poa, Agrostis,<br />
Phleum, Alopecurus, Phalaris and, <strong>to</strong> a lesser extent, in Holcus.<br />
Clusiidae<br />
(Figs: larva, 578, puparia 1202-1205)<br />
Over 200 species <strong>of</strong> Clusiidae have been described from most regions <strong>of</strong> <strong>the</strong> world.<br />
Ten species representing 4 genera occur in <strong>the</strong> <strong>British</strong> Isles. Little is known <strong>of</strong> <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> which all appear <strong>to</strong> develop in rotting wood. The larvae are said <strong>to</strong> be<br />
able <strong>to</strong> skip in a manner similar <strong>to</strong> Piophilidae.<br />
Clusiaflava (Meigen) has been reared from a dry beech stump (Smith, 1950). There<br />
are also two puparia in <strong>the</strong> <strong>British</strong> Museum (Nat. Hist.) from a rotten birch stump.<br />
Unfortunately, <strong>the</strong> caps containing <strong>the</strong> mouthparts have not survived from any <strong>of</strong> <strong>the</strong><br />
puparia available so I am unable <strong>to</strong> figure <strong>the</strong>m. The characteristic 'horned' puparium<br />
is illustrated (fig. 1202).<br />
Clusiodes albimana (Meigen) (figs 1203, 1205) has been reared several times from old<br />
logs and mossy tree stumps but in <strong>the</strong> puparia examined <strong>the</strong> ventral part <strong>of</strong> <strong>the</strong> cap, with<br />
<strong>the</strong> mouth parts, has been missing.<br />
There is a specimen <strong>of</strong> Clusiodes gentilis (Collin) with a puparium, 'bred from a log'<br />
by H. Oldroyd in <strong>the</strong> <strong>British</strong> Museum (Nat. Hist.), but <strong>the</strong> half <strong>of</strong> <strong>the</strong> puparial cap with<br />
<strong>the</strong> mouth parts is missing. However, Se guy ( 1934) gives an outline <strong>of</strong> <strong>the</strong> mouthparts<br />
which are illustrated here (fig. 578). Also in <strong>the</strong> BM (NH) collections is a specimen <strong>of</strong><br />
Clusiodes ( Columbiella) verticalis (Collin) with a puparium (fig. 1204), from 'old grass<br />
heap, back cottage garden'.<br />
Odiniidae<br />
(Figs: larvae 579-585, puparium 1206)<br />
Some 63 species in <strong>10</strong> genera <strong>of</strong> Odiniidae are known throughout <strong>the</strong> world. Of<br />
<strong>the</strong>se <strong>the</strong> larval habits <strong>of</strong> only I 0 species are known and all are associated with<br />
wounded, dead or dying trees. These have mostly been associated with <strong>the</strong> galleries<br />
<strong>of</strong> wood boring beetles (Buprestidae, Curculionidae, Scolytidae, Tenebrionidae,<br />
Oedemeridae), but some occur in <strong>the</strong> galleries <strong>of</strong> Diptera larvae (Hesperinidae,<br />
Pachyneuridae) and Lepidoptera (Cossidae, Gelechiidae). Most are considered <strong>to</strong> be<br />
saprophagous, or possibly feed on frass or dead insects. <strong>An</strong> American species (Odinia<br />
conspicua Sabrosky) is apparently predaceous on Tenebrionidae (Col.) in decayed<br />
poplar. The Asian species Turanodinia coccidarum Stackelberg was reported <strong>to</strong> have<br />
been reared from egg-masses <strong>of</strong> Pseudococcus coms<strong>to</strong>cki Kuwana.<br />
Seven species occur in Britain, all belonging <strong>to</strong> <strong>the</strong> genus Odinia, four <strong>of</strong> which have<br />
been reared.<br />
The larva <strong>of</strong> Odinia xanthocera Collin (figs 582- 585), described by V os de Wilde<br />
( 1935), is associated in Europe and North America with <strong>the</strong> weevil Cryp<strong>to</strong>rhynchus<br />
lapathi (L.), which mines twigs <strong>of</strong> Alnus, Salix and Populus. Odinia meijerei Collin (figs<br />
579- 581) is found on diseased elm trees in <strong>the</strong> tunnels <strong>of</strong> Scolytus (Lewis, 1979). Larvae<br />
<strong>of</strong> 0. pomona Cogan have been found under bark on apple. Odinia ornata Zetterstedt<br />
<strong>10</strong>3
occurs in beetle galleries on spruce in <strong>the</strong> USSR. Odinia hendeli Collin occurs in elm<br />
with larvae <strong>of</strong> Oedomeridae (Col.). Odinia maculata (Meigen) has been found in oak<br />
infested with Cossus (goat moth) larvae.<br />
Krivosheina ( 1981) reviews <strong>the</strong> systematics and biology <strong>of</strong> <strong>the</strong> Palaearctic species.<br />
Carnidae<br />
(Figs: larvae 586-589, egg <strong>10</strong>47, puparia 1207- 1208)<br />
Some 70 species <strong>of</strong> Carnidae have so far been described and include some <strong>of</strong> <strong>the</strong><br />
smallest Acalyptrates. They are mainly Palaearctic or Nearctic, some Holarctic in<br />
distribution with very few occurring in o<strong>the</strong>r regions. The larvae live in birds' nests<br />
where <strong>the</strong>y are saprophagous <strong>to</strong> semi-parasitic (Carnus, I <strong>British</strong> species) or<br />
saprophagous (Meoneura, 12 <strong>British</strong> species). Very little is known <strong>of</strong> <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong>.<br />
The larvae <strong>of</strong> Carnus hemapterus Nitzsch are found in <strong>the</strong> nest debris <strong>of</strong> a number <strong>of</strong><br />
birds including starling, hedge-sparrow, blackbird, kestrel and great spotted woodpecker.<br />
The tiny adults are blood sucking parasites <strong>of</strong> birds and break <strong>of</strong>f <strong>the</strong>ir own<br />
wings after settling down in <strong>the</strong> plumage <strong>of</strong> nestling birds. De Meijere (1912a) and<br />
Engel (1930) give partial descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> (figs <strong>10</strong>47, 1207).<br />
Meoneura species are associated with birds' nests as follows: M. lamellata Collin<br />
(crow, great tit, magpie, sand-martin, starling, 'thrush', etc.); M. neottiophila Collin<br />
(kestrel, sparrow-hawk, s<strong>to</strong>ck-dove, wren); M . obscurella (Fallen) (blackbird,<br />
sand-martin, 'finch', 'thrush'); M. lacteipennis (Fallen) and M. vagans (Fallen)<br />
(sand-martin); M. seducta Collin (herring-gull).<br />
Meoneura species are also believed <strong>to</strong> develop in animal excrement and carrion on<br />
which adults are sometimes found. In France M. exigua Collin has been reared from a<br />
damaged puparium <strong>of</strong> Sarcophaga.<br />
Acar<strong>to</strong>phthalmidae<br />
Only three species in one genus (Acar<strong>to</strong>phthalmus) <strong>of</strong> this family are known, two<br />
Holarctic and one Palaearctic. The two Holarctic species occur in Britain. Adults have<br />
been found on decaying fungi, dung and carrion. Ozerov ( 1987) found eggs on carrion<br />
and reared larvae on meat in <strong>the</strong> labora<strong>to</strong>ry.<br />
Periscelididae<br />
(Fig. 593, larva)<br />
About 20 species <strong>of</strong> Periscelididae are known from <strong>the</strong> Nearctic, Palaearctic and<br />
Neotropical regions. Four species <strong>of</strong> Periscelis occur in Britain.<br />
The life-his<strong>to</strong>ries and <strong>immature</strong> <strong>stages</strong> were unknown until very recently, when it was<br />
established that <strong>the</strong> larvae develop in <strong>the</strong> sap <strong>of</strong> tree wounds. The larva superficially<br />
resembles a Fannia (Fanniidae) and is illustrated (fig. 593) here from a figure appearing<br />
in Teskey ( 1976). Dr Tony lrwin has found larvae <strong>of</strong> Microperiscelis annulata (Fallen)<br />
in ash seepage in Norwich.<br />
Aulacigastridae<br />
(Figs: larva 590-591, egg <strong>10</strong>48, puparium, 12<strong>10</strong>)<br />
This is a small and little-known family <strong>of</strong> about 4 species in 3 genera represented in<br />
<strong>the</strong> major zoogeographical regions except <strong>the</strong> Australian and Pacific. Only one species<br />
occurs in Britain.<br />
<strong>10</strong>4
The larvae <strong>of</strong> Aulacigaster /eucopeza (Meigen) develop in sap fluxes exuded from<br />
tree wounds. Robinson, I. ( 1953) describes <strong>the</strong> <strong>immature</strong> <strong>stages</strong>. At pupation <strong>the</strong><br />
prothoracic spiracles are completely everted and protrude from <strong>the</strong> surface <strong>of</strong> <strong>the</strong> slime<br />
flux in which <strong>the</strong> puparium is submerged.<br />
Stenomicridae<br />
(Figs: larva 592, egg <strong>10</strong>49, puparium 1209)<br />
The problematic and little-known genus Stenomicra has recently been given family<br />
status in <strong>the</strong> Palaearctic Catalogue. The genus contains some 20 described species and<br />
is represented in all <strong>the</strong> major zoogeographical regions. Only one <strong>British</strong> species was<br />
known until Irwin (1982) described a second from Wales. Irwin also discusses <strong>the</strong><br />
taxonomic peregrinations <strong>of</strong> <strong>the</strong> genus. Nothing is known <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ry or<br />
<strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> five Palaearctic species. Williams (1939) records larvae in<br />
water-holding leaf bases <strong>of</strong> Gramineae (Poaceae) and o<strong>the</strong>r plants in Hawaii (fig. 592).<br />
<strong>An</strong>thomyzidae<br />
(Figs: larvae 594-606)<br />
The family <strong>An</strong>thomyzidae is represented in all zoogeographical regions by a <strong>to</strong>tal <strong>of</strong><br />
about 60 species. Sixteen species in 4 genera occur in Britain.<br />
The larvae develop between <strong>the</strong> closely fitting leaf sheaths <strong>of</strong> water-loving<br />
Gramineae and occur in <strong>the</strong> galls formed by Lipara (Chloropidae). One species has<br />
been reared from fungi.<br />
The larvae <strong>of</strong> <strong>An</strong>thomyza ( = Stiphrosoma) sabulosa (Haliday) (figs 594-597) feed<br />
between <strong>the</strong> leaf sheaths <strong>of</strong> tillers <strong>of</strong> Arrhena<strong>the</strong>rum e/atius L. and Lolium perenne L.,<br />
but do not sever <strong>the</strong> central shoot and cause little damage. They overwinter in <strong>the</strong> third<br />
instar and pupate <strong>to</strong>wards <strong>the</strong> end <strong>of</strong> April. Third ins tar larvae have also been found in<br />
July so <strong>the</strong>re are probably two generations per year. The larvae <strong>of</strong> A. gracilis Fallen<br />
(figs 598-602), A. sordide/la (Zetterstedt) and A. bifasciata Wood have been found<br />
between <strong>the</strong> leaves <strong>of</strong> Phragmites and Typha. <strong>An</strong>thomyza a/bimana (Meigen) has been<br />
reared from fungi (Agaricales). O<strong>the</strong>r species have been reared from birds' nests<br />
(presumably from grasses used in <strong>the</strong>ir construction), e.g. A . cingu/ata Haliday (rook)<br />
and A . neg/ecta Collin (coot) (Collin, 1944).<br />
Paranthomyza nitida (Meigen) (figs 603-606) has been recorded from Melandrium<br />
rubrum (Wiegel) (de Meijere 1944).<br />
<strong>An</strong>agnota bicolor (Meigen) has been reared from a mole's nest (Collin, 1944),<br />
presumably from grasses used in nesting material.<br />
Asteiidae<br />
Some 130 species in <strong>10</strong> genera <strong>of</strong> Asteiidae have been described and <strong>the</strong> family is<br />
represented in all <strong>of</strong> <strong>the</strong> major zoogeographical regions. Seven species in 3 genera occur<br />
in Britain (Chandler, 1978). The <strong>immature</strong> <strong>stages</strong> have not been described.<br />
Asteia amoena Meigen is said <strong>to</strong> develop in wood detritus, in hollow trees, etc.,<br />
but A. concinna Meigen adults are associated with marshes, fens, dune slacks and<br />
Phragmites beds.<br />
Adults <strong>of</strong> Astiosoma rufifrons Duda are attracted <strong>to</strong> cold wood ash and smouldering<br />
bonfires <strong>of</strong> elm branches, etc., but <strong>the</strong> life-his<strong>to</strong>ry is unknown.<br />
Leiomyza species have been reared from fungi: L. dudai Sabrosky and L. sca<strong>to</strong>phagina<br />
(Fallen) (several agarics, chiefly lignicolous, and <strong>the</strong> latter also Po/yporus squamosus<br />
<strong>10</strong>5
Fries); L. laevigata (Meigen) (Pleurotus cornucopiae L.). Larvae <strong>of</strong> L. sca<strong>to</strong>phagina<br />
have also been recorded from dried stems <strong>of</strong> <strong>the</strong> common reed (Phragmites) <strong>to</strong>ge<strong>the</strong>r<br />
with Lipara (Chloropidae) (Colyer & Hammond, 1968).<br />
Camillidae<br />
Eleven species <strong>of</strong> Camillidae are known, all contained in <strong>the</strong> genus Camilla and<br />
represented in only <strong>the</strong> Palaearctic (9), Nearctic (I) and Afrotropical regions (3).<br />
No <strong>immature</strong> <strong>stages</strong> <strong>of</strong> this family have been described. Collin ( 1933) notes that E. B.<br />
Basden found adults <strong>of</strong> C. fuscipes Coli in at <strong>the</strong> entrance <strong>of</strong> rabbit burrows and has<br />
reared C. a tripes Duda (as acutipennis Loew) from nests <strong>of</strong> <strong>the</strong> short-tailed field vole.<br />
Baumann ( 1977) found adults <strong>of</strong> C. atripes (as C. atrimana Strobl) in <strong>the</strong> nests <strong>of</strong> voles<br />
in Germany. Dr A. G . lrwin (pers. comm.) has reared a species from soil derived from<br />
feral pigeon guano.<br />
Ephydridae<br />
(Figs: larvae 6<strong>10</strong>--624, eggs I 050--<strong>10</strong>56, puparia 1211- 1218, leaf-mine 1304)<br />
Over I ,500 species <strong>of</strong> Ephydridae (shore flies) are known from all <strong>the</strong> major<br />
zoogeographical regions. Some 130 species in 39 genera and 4 subfamilies occur in<br />
Britain.<br />
The larvae are mainly aquatic or semi-aquatic and over <strong>the</strong> world are found in a very<br />
wide range <strong>of</strong> habitats including some <strong>of</strong> <strong>the</strong> harshest in which Diptera are known <strong>to</strong><br />
develop, e.g. hot springs (some Ephydra, Scatella, Paracoenia), petroleum pools<br />
(Halaeomyia), brine (Ephydra), egg-pods <strong>of</strong> locusts (Ac<strong>to</strong>ce<strong>to</strong>r), scavengers in terrestrial<br />
snail shells (Discomyza), in spider egg-cases (Trimerina), in carrion and faeces<br />
(Allotrichoma, Hecamede), in nests <strong>of</strong> ants (Rhyncopsilopa) and birds (Nostima), as<br />
serious pests <strong>of</strong> rice (Hydrellia) and <strong>of</strong> beet (Clanoneurum).<br />
All <strong>stages</strong> may be present (adults 'graze' algae on mud surfaces, s<strong>to</strong>nes, etc.) in very<br />
large numbers and may thus provide an important food source for marsh and shore<br />
frequenting birds and even man (American Indians).<br />
In spite <strong>of</strong> all this <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and ecology <strong>of</strong> <strong>the</strong>se highly successful<br />
and adaptable flies remain poorly known. R. Dahl (1959) provides a pioneer ecological<br />
study for Scandinavia. A valuable symposium volume on Ephydridae (Deonier, 1979)<br />
includes much biological and ecological information.<br />
The larvae <strong>of</strong> only 25 <strong>of</strong> <strong>the</strong> <strong>British</strong> species in 14 genera have been described, mostly<br />
by continental or American authors. The early literature on <strong>the</strong> <strong>immature</strong> <strong>stages</strong> is<br />
summarised, with illustrations, by R. Dahl (1969) and Hennig (1943a). O<strong>the</strong>r useful<br />
works are listed below. Genera not mentioned below may be assumed <strong>to</strong> be unknown in<br />
<strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Psilopinae. Psilopa leucos<strong>to</strong>ma (Meigen) has been reared from mines in Chenopodium<br />
(Oidroyd, 1964); <strong>the</strong> larva <strong>of</strong> <strong>the</strong> Nearctic P. petrolei Coquillett (now referred <strong>to</strong><br />
Halaeomyia) is unique in that it lives, swims and feeds (on <strong>the</strong> bodies <strong>of</strong> trapped insects)<br />
in pools <strong>of</strong> crude petroleum in California (Thorpe, 1930).<br />
Discocerina obscure/la (Fallen) (figs 611, 612, I 050, 1211) probably feeds on microorganisms<br />
(Foote & Eastin, 1974).<br />
Discomyza incurva (Fallen) (fig. 6<strong>10</strong>) has been reared from a decaying snail (Helix<br />
pomatia L.).<br />
Larvae <strong>of</strong> Clanoneurum cimiciforme (Haliday) mine Salicornia stricta Dum<br />
(glasswort).<br />
<strong>10</strong>6
The larvae <strong>of</strong> Trimerina madizans (Fallen) feed as parasi<strong>to</strong>ids within <strong>the</strong> egg cases <strong>of</strong><br />
marsh-dwelling spiders (Miryphantes).<br />
Notiphilinae. Notiphila brunnipes Robineau-Desvoidy lays eggs on <strong>the</strong> flowers <strong>of</strong> <strong>the</strong><br />
white water-lily. The flowers close at sunset and withdraw below <strong>the</strong> water surface,<br />
reappearing at sunrise. The eggs are thus alternately immersed and exposed. The larvae<br />
live in <strong>the</strong> stems. The larvae (fig. 615) <strong>of</strong> most species <strong>of</strong> No<strong>to</strong>phila however live in <strong>the</strong><br />
soil at <strong>the</strong> bot<strong>to</strong>m <strong>of</strong> lakes, ponds and streams and obtain <strong>the</strong>ir oxygen from <strong>the</strong><br />
intercellular spaces in <strong>the</strong> roots <strong>of</strong> aquatic plants by means <strong>of</strong> sharp hollow terminal<br />
spines (figs 616, 1213) (see Varley, 1937; Keilin, 1944; Houlihan, 1969; and compare<br />
Chrysogaster, Syrphidae; Erioptera, Tipulidae).<br />
All Hydrellia larvae are phy<strong>to</strong>phagous. Most species mine <strong>the</strong> leaves and stems <strong>of</strong><br />
aquatic plants but o<strong>the</strong>rs attack terrestrial plants, especially grasses (including rice). H.<br />
chrysos<strong>to</strong>ma (Meigen) lives on Potamoge<strong>to</strong>n as do many o<strong>the</strong>r European and Nearctic<br />
species. H. nasturtii Collin (figs 613, 1304) mines <strong>the</strong> stems <strong>of</strong> watercress (Taylor, 1928)<br />
and can be <strong>of</strong> economic importance (compare Scap<strong>to</strong>myza, Drosophilidae). The larva<br />
<strong>of</strong> H. modesta Loew (fig. 614) mines and overwinters in <strong>the</strong> basal part <strong>of</strong> <strong>the</strong> leaf blade<br />
and leaf sheath <strong>of</strong> grasses (Holcus, <strong>An</strong>thoxanthum) but does not penetrate <strong>to</strong> <strong>the</strong> central<br />
shoot (Nye, 1958). H. griseola (Fallen) (some records <strong>of</strong> which may refer <strong>to</strong> <strong>the</strong> previous<br />
species) does similar damage <strong>to</strong> cereals. Attacked leaves turn yellow, starting at <strong>the</strong> tip,<br />
and wi<strong>the</strong>r away. H. griseola, when very common in any locality, is said <strong>to</strong> break its<br />
normal habit <strong>of</strong> mining in only monocotyledonous plants by mining in dicotyledons <strong>of</strong><br />
considerable diversity, e.g. Lychnis, Trifolium and many o<strong>the</strong>rs (Hering, 1951 ).<br />
Parydrinae. The larva <strong>of</strong> Parydra pus ilia (Meigen) was found in wet moss and in mud<br />
at <strong>the</strong> bot<strong>to</strong>m <strong>of</strong> a lake in Iceland (Nielsen, P. et al., 1954).<br />
Vim mer (1925) briefly describes <strong>the</strong> larva <strong>of</strong> Pelina aenescens Stenhammer found in a<br />
quiet stream.<br />
First and second instar larvae <strong>of</strong> <strong>the</strong> Holarctic Och<strong>the</strong>ra mantis (De Geer) (fig. 618)<br />
are obligate preda<strong>to</strong>rs <strong>of</strong> larval Chironomidae but third instars will feed on o<strong>the</strong>r<br />
aquatic insect larvae and can be cannibalistic in <strong>the</strong> absence <strong>of</strong> prey (Simpson, 1975).<br />
Three species are now known <strong>to</strong> occur in Britain (lrwin, 1985b).<br />
Ephydrinae. The larva <strong>of</strong> Ephydra riparia Fallen has well developed abdominal<br />
prolegs with claw-like spines (crochets) an adaptation for life on aquatic plants beneath<br />
<strong>the</strong> water surface in saltmarshes. The puparium is adapted for attaching itself <strong>to</strong> plants<br />
by a curious excision <strong>of</strong> <strong>the</strong> posterior segments; <strong>the</strong> posterior spiracles are situated at<br />
<strong>the</strong> ends <strong>of</strong> <strong>the</strong> branches <strong>of</strong> a long forked posterior appendage (fig. 1214). Beyer (1939)<br />
describes <strong>the</strong> larvae <strong>of</strong> E. riparia, Setacera micans (Haliday) (fig. 617) and Paracoenia<br />
.fumosa (Stenhammer). The larva <strong>of</strong> <strong>the</strong> abundant Scatella stagna/is (Fallen) feeds on a ·<br />
wide variety <strong>of</strong> algae. S. silacea Loew (figs 619- 621) breeds in sewage filter beds (Terry,<br />
1952). According <strong>to</strong> Bollwig ( 1940) <strong>the</strong> larvae <strong>of</strong> Sca<strong>to</strong>phila unicorn is Czerny cover<br />
<strong>the</strong>ir bodies with small s<strong>to</strong>nes or faecal pellets.<br />
Teichomyza fusca Macquart, because <strong>of</strong> its unsavoury habits, now enjoys a<br />
cosmopolitan distribution. The fly appears wherever <strong>the</strong>re are cesspits, urine soaked<br />
woodwork or o<strong>the</strong>r materials, or cadavers at <strong>the</strong> stage when nutritious liquids ooze<br />
from collapsing tissues. The larvae (figs 622-624) are very gregarious and may assemble<br />
in such masses as <strong>to</strong> block drains and septic tanks. Sometimes larvae are swallowed in<br />
foul water and have been reported in cases <strong>of</strong> urinary and rectal myiasis (Zumpt, 1965).<br />
The puparia (fig. 1218) are <strong>of</strong>ten encountered in numbers during archaeological<br />
excavations and indicate <strong>the</strong> location <strong>of</strong>middens and cesspits.<br />
Oldroyd (1964) aptly summarized <strong>the</strong> biological versatility <strong>of</strong> this family<br />
" Evidently we are seeing in <strong>the</strong> Ephydridae a family <strong>of</strong> flies in <strong>the</strong> full flower <strong>of</strong> its<br />
<strong>10</strong>7
evolution, and as such <strong>the</strong>y <strong>of</strong>fer attractive material for study, not only <strong>to</strong> <strong>the</strong> dipterist,<br />
but also <strong>to</strong> students <strong>of</strong> insect physiology and behaviour".<br />
Diastatidae<br />
(Figs: larva 607- 609)<br />
Some 33 species <strong>of</strong>Diastatidae are known from <strong>the</strong> major zoo-geographical regions<br />
(except <strong>the</strong> Australo-Pacific). In Britain two species <strong>of</strong> Campichoeta and six <strong>of</strong> Diastata<br />
occur. Almost nothing is known <strong>of</strong> <strong>the</strong>ir life-his<strong>to</strong>ries, but <strong>the</strong>y are probably spent in<br />
decaying wood. Hennig (1952) illustrates what he believes <strong>to</strong> be Campichoeta basalis<br />
(Meigen) (=punctum (Meigen)) (figs 607- 609).<br />
Drosophilidae<br />
(Figs: larvae 625-638, eggs <strong>10</strong>57- <strong>10</strong>63, puparia 1219- 1229, leaf-mines 1296-1298)<br />
Over 2,500 species <strong>of</strong> Drosophilidae have so far been described throughout <strong>the</strong><br />
world. The particular association <strong>of</strong> <strong>the</strong> family with decaying and fermenting fruits and<br />
vegetables, wine, beer, etc. have earned <strong>the</strong>m several common names, e.g. fruit-flies,<br />
pomace-flies, vinegar-flies, etc. Lesser fruit-flies has been used <strong>to</strong> distinguish<br />
Drosophilidae from Tephritidae, <strong>the</strong> true fruit-flies. This association has also ensured a<br />
cosmopolitan distribution via commerce for several species. O<strong>the</strong>r substances in which<br />
larvae develop include fungi, slime fluxes, dung, carrion and many are general scavengers.<br />
Some mine <strong>the</strong> leaves and stems <strong>of</strong> plants, some are symbionts <strong>of</strong> bees, and o<strong>the</strong>rs<br />
are preda<strong>to</strong>rs <strong>of</strong> small Homoptera.<br />
Because <strong>of</strong> <strong>the</strong>ir rapid rate <strong>of</strong> development and ease <strong>of</strong> culture on artificial media<br />
many Drosophila species (e.g. D. melanogaster Meigen) have become important labora<strong>to</strong>ry<br />
animals in research on genetics, cy<strong>to</strong>logy, physiology and ethology. Consequent<br />
upon this <strong>the</strong> <strong>immature</strong> <strong>stages</strong> and labora<strong>to</strong>ry life-his<strong>to</strong>ries <strong>of</strong> many species have<br />
become well known. However, our knowledge <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ries and ecology <strong>of</strong><br />
Drosophilidae in field conditions is not extensive, and <strong>the</strong>re is much scope for research.<br />
The adults are attracted <strong>to</strong> a far greater range <strong>of</strong> apparently suitable breeding media<br />
than that which actually proves suitable for development.<br />
Okada (1968) gives a very useful account <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> family and<br />
Ashburner (1981) reviews <strong>the</strong> life-his<strong>to</strong>ries <strong>of</strong> 'bizarre' Drosophilidae. Shorrocks<br />
(1972) provides a valuable <strong>introduction</strong> <strong>to</strong> all aspects <strong>of</strong> <strong>the</strong> biology <strong>of</strong> <strong>the</strong> genus<br />
Drosophila and Basden (1954) is useful. Labora<strong>to</strong>ry workers will find <strong>the</strong> Drosophila<br />
Information Service (DIS) (Department <strong>of</strong> Biology, University <strong>of</strong> Oregon, Eugene,<br />
Oregon, USA) invaluable for annual updating <strong>of</strong> knowledge on Drosophila mutants,<br />
species maintained in cultures throughout <strong>the</strong> world, research notes and techniques in<br />
culturing.<br />
Some 52 species in I 0 genera and two subfamilies occur in Britain. The classification<br />
below follows Biichli & Pite (in So6s & Papp, 1984), which differs from <strong>the</strong> Handbook<br />
Check List. Genera and species not mentioned may be assumed <strong>to</strong> be unknown in <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong>.<br />
Steganinae. The larva <strong>of</strong> Steganina coleoptrata (Scopoli) (figs 625-627) feeds on sap<br />
under <strong>the</strong> bark <strong>of</strong> birch and Prunus (Morge, 1956) and has also been reared from fungi<br />
(Hypoxylon fragiforme). Ac/e<strong>to</strong>xenus formosus (Loew) is predaceous on Siphoninus<br />
immaculatus (Heeger) (on ivy) in Britain and o<strong>the</strong>r species <strong>of</strong> Aleyrodidae (Homoptera)<br />
(on hawthorn and Viburnum) abroad. The larva is orange at first becoming green in <strong>the</strong><br />
third ins tar and is covered in a mucilagenous slime which collects debris, including eggshells<br />
<strong>of</strong> <strong>the</strong>ir whitefly prey. The puparia (fig. 1220) are whitish and flattened ventrally<br />
where <strong>the</strong>y are firmly glued <strong>to</strong> <strong>the</strong> old ivy (Hedera helix) leaves on which <strong>the</strong>y occur. The<br />
<strong>10</strong>8
Acle<strong>to</strong>xenus larva may consume 30--40 whitefiy 'puparia' during its development but<br />
attempts at biological control against whitefiy have not been successful (Ashburner,<br />
1981;Ciausen&Berry, 1932).<br />
Cacoxenus indaga<strong>to</strong>r Loew (figs 628-631) is a nest parasite <strong>of</strong> many species <strong>of</strong><br />
solitary bees (<strong>An</strong>thophora, Chalicodoma, Osmia) (Julliard, 1947; de Meijere, 1944).<br />
The larva eats supplies provided for <strong>the</strong> bee progeny. It appears that both bee and<br />
drosophilid larva can survive in <strong>the</strong> same cell if enough food is provided, but if not <strong>the</strong><br />
Cacoxenus may eat <strong>the</strong> host larva, possibly after it dies <strong>of</strong> starvation. Careful research is<br />
needed on <strong>the</strong> detail <strong>of</strong> this life-his<strong>to</strong>ry. I have had larvae <strong>of</strong> Cacoxenus sent in <strong>to</strong> me<br />
from behind door knobs in a school; clearly solitary bees had been nesting in <strong>the</strong> old<br />
style mortice locks. Larvae have also been found in sash window-frames along with bee<br />
(Osmia) larvae and Ptinus beetles.<br />
Amiota ( Amiota) alboguttata (Wahlberg) has been reared from fungi (Daldinia<br />
concentrica, Pyrenomycetae). Abroad, some Amiota species are predaceous on<br />
Homoptera (Pseudococcidae, Delphacidae) on sugarcane, and one species is found in<br />
<strong>the</strong> burrows <strong>of</strong> an ambrosia-beetle pest on tea where it may feed on fungi with which <strong>the</strong><br />
beetle lives in symbiotic association (Ash burner, 1981). Amiota ( Phortica) variegata<br />
(Fallen) (fig. 632) larvae feed on sap bleeding from oak, chestnut and willow. The three<br />
<strong>British</strong> species, as yet unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>, may <strong>the</strong>refore be found <strong>to</strong> have<br />
unusual or interesting life-his<strong>to</strong>ries.<br />
Leucophenga maculata (Dufour) has been reared from several fungi (agarics, boleti<br />
and polypores).<br />
Drosophilinae. Scap<strong>to</strong>myza species (figs 633-634, 1296--1298) are leaf-miners in <strong>the</strong><br />
following plants: S. flava (Fallen) (Cruciferae (Brassicaceae), including Brassica);<br />
S. graminum (Fallen) (A trip/ex, Brassica, Chenopodium, Lychnis, Melandrium, Silene,<br />
Stellaria, Viscaria, also stems <strong>of</strong> watercress): S. pallida (Zetterstedt) (clover) and<br />
S. disticha Duda (Allium).<br />
Many species <strong>of</strong> Drosophila (figs 635-638) develop in rotting fruit and vegetable<br />
matter and fermenting substances. D. rep/eta Wollas<strong>to</strong>n is a cosmopolitan species that<br />
has become a nuisance in restaurants, kitchens, canteens and taverns, where it breeds<br />
in nearby decomposing vegetables, including onions and cabbages. The common<br />
ferment-loving species include D. (D.) funebris (F.), D.fenestratum Fallen, D.fasciata<br />
Meigen and D. obscura Fallen. Species reared from fungi include D. busckii Coquillett,<br />
D. cameraria Haliday, D. confusa Staeger, D.funebris (F.), D. his trio Meigen, D. kuntzei<br />
Duda, D. melanogaster Meigen, D. phalerata Meigen, D. obscura Fallen, D. rep/eta<br />
Wollas<strong>to</strong>n, D. subobscura Collin and D. transversa Fallen. Chandler (in Stubbs &<br />
Chandler, 1978) lists <strong>the</strong> species <strong>of</strong> host fungi. Drosophila reared from sap exudates<br />
on trees include D. lit<strong>to</strong>ralis Meigen, D. obscura and D. subobscura. D. ananassae<br />
Doleschall and D .funebris have been reared from excrement. D. busckii has been reared<br />
from carrion.<br />
Puparia <strong>of</strong> D. busckii (fig. 1225) and D.funebris (fig. 1226) are frequently found in<br />
milk bottles where <strong>the</strong>y are so firmly stuck <strong>to</strong> <strong>the</strong> glass <strong>the</strong>y come through <strong>the</strong> washing<br />
process intact (though killed by it). Some may float <strong>of</strong>f in<strong>to</strong> <strong>the</strong> milk when <strong>the</strong> bottles<br />
are refilled and be brought <strong>to</strong> <strong>the</strong> notice <strong>of</strong> environmental health <strong>of</strong>ficers (see also<br />
Spiniphora bergenstammi, Phoridae).<br />
The puparia and eggs <strong>of</strong> Drosophila species are very distinctive because <strong>of</strong> <strong>the</strong>ir long<br />
respira<strong>to</strong>ry processes and a range <strong>of</strong> commonly encountered species is shown (figs<br />
<strong>10</strong>59-<strong>10</strong>63, 1225-1229).<br />
A key <strong>to</strong> drosophilid larvae is provided by Okada (1968) but because <strong>of</strong> <strong>the</strong> rapid<br />
development rate it is frequently more convenient <strong>to</strong> rear living larvae through <strong>to</strong><br />
adults for identification purposes using <strong>the</strong> keys <strong>of</strong> Fonseca ( 1965) or Basden ( 1954).<br />
Shorrocks ( 1972) provides a key <strong>to</strong> adults <strong>of</strong> <strong>the</strong> common species <strong>of</strong> Drosophila.<br />
<strong>10</strong>9
Milichiidae<br />
(Figs: larvae 639-644, puparia 1230-1231)<br />
Some 245 species <strong>of</strong> Milichiidae are known throughout <strong>the</strong> world. The larvae are<br />
coprophagous or saprophagous and have been reared from excrement, kitchen refuse,<br />
plant detritus, birds' nests and ants' nests. Eleven species in 5 genera and 2 subfamiles<br />
occur in Britain.<br />
Madazinae. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> two <strong>British</strong> species <strong>of</strong> Madiza are unknown<br />
but adults <strong>of</strong> M . glabra Fallen are associated with primitive sanitary arrangements in<br />
country areas and sometimes occur in large numbers indoors. Adults have also been<br />
found on cadavers during 'caseic' or protein fermentation especially if putrid liquids<br />
exude.<br />
Lep<strong>to</strong>me<strong>to</strong>pa and Desmome<strong>to</strong>pa are known <strong>to</strong> breed in human excrement. Hennig<br />
( 1956) has described <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Lep<strong>to</strong>me<strong>to</strong>pa (figs 639- 644).<br />
Phyllomyza formicae Schmitz and P. securicornis Fallen have been found in ants'<br />
nests (Formica rufa L.) (Donisthorpe, 1927) and Hennig (1937) figures <strong>the</strong> puparia<br />
(figs 1230-1231 ).<br />
Milichiinae. The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Milichia ludens (Wahlberg) have not been<br />
described but <strong>the</strong> species is associated with <strong>the</strong> ant Lasius fuliginosus (Latreille)<br />
according <strong>to</strong> Donisthorpe (1927), who fur<strong>the</strong>r states that <strong>the</strong> larvae <strong>of</strong> an African<br />
species covers itself with excreta and wanders about in ant runs.<br />
The larva <strong>of</strong> an American milichiid feeds on exhausted fungus garden substrates and<br />
refuse in ants' nests (Moser & Neff, 1971).<br />
Tethinidae<br />
Ninety-two species <strong>of</strong> Tethinidae are known throughout <strong>the</strong> world, <strong>of</strong> which <strong>10</strong><br />
species in 2 genera occur in Britain.<br />
The <strong>immature</strong> <strong>stages</strong> and life-his<strong>to</strong>ries are unknown but <strong>the</strong> adults are found in<br />
coastal regions, on fore-shores or dunes, and in saline meadows and lagoons.<br />
Canacidae<br />
(Figs: larvae 645-646, puparia 1233)<br />
Ninety-one species <strong>of</strong>Canacidae are known throughout <strong>the</strong> world. Two species in 2<br />
genera occur in Britain.<br />
The larva <strong>of</strong> Canace nasica Haliday feeds on Enteromorpha (Algae) in <strong>the</strong> coastal<br />
intertidal zone. The puparium (fig. 1233) is described by Hin<strong>to</strong>n (1967a) who established<br />
its possession <strong>of</strong> plastron bearing spiracular gills in common with a number <strong>of</strong><br />
o<strong>the</strong>r intertidal Diptera. The larval <strong>stages</strong> <strong>of</strong> an American Canace (figs 645-646) have<br />
also been found <strong>to</strong> feed on blue-green algae in <strong>the</strong> intertidal zone (Teskey & Valiela,<br />
1977). The <strong>immature</strong> <strong>stages</strong> and life-his<strong>to</strong>ry <strong>of</strong> Xanthocanace ranula (Loew) are<br />
unknown.<br />
Braulidae<br />
(Figs: larvae 64 7-649, egg <strong>10</strong>64, puparia 1232)<br />
Only 8 species <strong>of</strong> Braulidae, contained in two genera, Braula and Megabraula, are<br />
known. The tiny, wingless mite-like adults feed on pollen in <strong>the</strong> nests <strong>of</strong> honey-bees,<br />
<strong>of</strong>ten attached <strong>to</strong> <strong>the</strong> bees. The larvae feed on pollen in <strong>the</strong> bees' brood combs. Not<br />
1<strong>10</strong>
surprisingly, <strong>the</strong> distribution <strong>of</strong> <strong>the</strong>se flies largely coincides with <strong>the</strong> distribution <strong>of</strong><br />
apiculture throughout <strong>the</strong> world.<br />
One species, Brau/a coeca Nitzsch (figs 647- 649), occurs in Britain, <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong> and life-his<strong>to</strong>ry <strong>of</strong> which have been described by several authors including Skaife<br />
(1921) and Imms (1942).<br />
Agromyzidae<br />
(Figs: larvae 650--668, eggs <strong>10</strong>65- <strong>10</strong>67, puparia 1234-1240, leaf-mines 1299- 1303)<br />
Over 2,000 species <strong>of</strong> Agromyzidae have been described throughout <strong>the</strong> world,<br />
about half that number occurring in <strong>the</strong> Palaearctic region. Some 320 species in 18<br />
genera and 2 subfamilies occur in Britain.<br />
The larvae feed on plant tissues, <strong>the</strong> majority (75%) as leaf miners, o<strong>the</strong>rs as stem<br />
borers, a few in roots, some developing in twigs on trees with some causing galls. About<br />
160 <strong>of</strong> <strong>the</strong> world species may have some economic importance (Spencer, 1973b).<br />
Most Agromyzidae are limited in <strong>the</strong>ir choice <strong>of</strong> host <strong>to</strong> a single plant species or,<br />
more <strong>of</strong>ten, several species within <strong>the</strong> same plant genus; o<strong>the</strong>rs feed on a number <strong>of</strong><br />
genera within a single plant family or on several related families <strong>of</strong> <strong>the</strong> same order; a few<br />
species are truly polyphagous on a wide range <strong>of</strong> unrelated hosts.<br />
The form <strong>of</strong> <strong>the</strong> mine can be <strong>of</strong>considerablehelp in <strong>the</strong> identification <strong>of</strong> Agromyzidae.<br />
The main types <strong>of</strong> mine are linear or serpentine (figs 1299, 1302- 1303) or form a blotch<br />
(fig. 1300). The part <strong>of</strong> <strong>the</strong> leaf that is mined may also be taxonomically important as<br />
may <strong>the</strong> arrangement <strong>of</strong>frass within <strong>the</strong> mine. It should also be borne in mind that some<br />
members <strong>of</strong> o<strong>the</strong>r families <strong>of</strong>Diptera also form mines (figs 1287- 1298, 1304-1308), and<br />
see under Sciaridae, Chironomidae, Dolichopodidae, Syrphidae, Scathophagidae,<br />
Tephritidae, Lauxaniidae, Psilidae, <strong>An</strong>thomyzidae, Drosophilidae, Ephydridae,<br />
Chloropidae and <strong>An</strong>thomyiidae.<br />
Eggs are laid on <strong>the</strong> plants, beneath <strong>the</strong> epidermis. Pupation normally takes place on<br />
<strong>the</strong> ground but a few species pupate at <strong>the</strong> end <strong>of</strong> <strong>the</strong> mine.<br />
Knowledge <strong>of</strong> <strong>the</strong> general biology <strong>of</strong> Agromyzidae is summarised by Hering (1951)<br />
and Spencer ( 1972, 1976). Spencer ( 1973b) deals with <strong>the</strong> species <strong>of</strong> economic importance.<br />
De Meijere (1925-1955) has described <strong>the</strong> larvae <strong>of</strong> many Agromyzidae and P.<br />
Alien ( 1956-1958) describes <strong>the</strong> larvae <strong>of</strong> several <strong>British</strong> species. Although <strong>the</strong> host<br />
plants <strong>of</strong> many species are known, detailed studies <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ries or morphology<br />
<strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are still few and <strong>the</strong>re is plenty <strong>of</strong> scope for fur<strong>the</strong>r research.<br />
To facilitate such research Griffiths (1962) provides useful information on rearing<br />
Agromyzidae and <strong>the</strong>ir parasites. Hering (1957) gives a useful account <strong>of</strong> mines, with<br />
keys.<br />
Third instar larvae <strong>of</strong> <strong>the</strong> two subfamilies may be distinguished as follows:<br />
Cephalopharyngeal skele<strong>to</strong>n with each dorsal cornu divided in<strong>to</strong> two (figs 651 --{)52) .<br />
. . . . . . . . . . . . . . . . Agromyzinae<br />
Dorsal cornua not divided (figs 654, 661-{)62, 668) . . . . . . . . Phy<strong>to</strong>myzinae<br />
Agromyzinae. The <strong>British</strong> species <strong>of</strong> Hex omyza all cause twig galls: H. simp/icoides<br />
(Hendel) on Sa/ix (especially S. caprea L.); H. sarothamni (Hendel) on Sarothamnus<br />
scoparius (L.) and H. schineri (Giraud) on Populus nigra L. and P. tremu/a L.<br />
Nine <strong>of</strong> <strong>the</strong> 13 <strong>British</strong> species <strong>of</strong> M elanagromyza (figs 650--651) are internal stemborers;<br />
M . cunctans (Meigen) forms slender stem galls on Lotus; M . symphyti Griffiths<br />
feeds in <strong>the</strong> thick leaf stalks <strong>of</strong> Symphytum <strong>of</strong>ficina/e L. The hosts <strong>of</strong> 4 species are<br />
unknown.<br />
Of <strong>the</strong> 23 <strong>British</strong> species <strong>of</strong> Ophiomyia (fig. 652) 13 feed as external stem-miners and<br />
one (0. me/andryi de Meijere) forms an internal stem mine. Five are leaf miners but<br />
111
mostly inside <strong>the</strong> midrib with only one species (0. maura (Meigen)) forming a typical<br />
mine in <strong>the</strong> leaf blade. The life his<strong>to</strong>ries <strong>of</strong> 4 species are unknown. Ophiomyia simplex<br />
(Loew) (<strong>the</strong> asparagus miner) is recorded as an occasional stem mining pest <strong>of</strong><br />
asparagus (H. F. Barnes, 1937; Spencer, 1973b) though <strong>the</strong> primary cause <strong>of</strong> damage<br />
may be a fungus (Spencer, 1976).<br />
Forty-three <strong>of</strong> <strong>the</strong> 46 <strong>British</strong> species <strong>of</strong> Agromyza are leaf miners (15 on Gramineae<br />
( = Poaceae) 12 on Papilionacae, <strong>the</strong> rest on various families) with one stem miner (A.<br />
marionae Griffiths), one stem gall causer (A . erythrocephala Hendel) (both on Vicia<br />
cracca L.) and one unknown (A . rubiginosa Griffiths). Several species <strong>of</strong> Agromyza<br />
attack cereals (fig. 1300) but rarely cause much damage in Britain, though <strong>the</strong>y can be<br />
serious pests abroad.<br />
Phy<strong>to</strong>myzinae. The larvae <strong>of</strong> all <strong>the</strong> Phy<strong>to</strong>bia species so far known feed on <strong>the</strong><br />
cambium <strong>of</strong> <strong>the</strong> twigs or trunks <strong>of</strong> trees. The larvae (fig. 656) can be as long as 2 cm,<br />
quite unlike <strong>the</strong> usual agromyzid, and may do permanent damage <strong>to</strong> <strong>the</strong> wood. P.<br />
cerasiferae (Kangas) has been found on Prunus cerasifera Ehrendorfer; P. carbonaria<br />
(Zetterstedt) is found on Rosaceae, Crataegus and Malus; P. cambii (Hendel) (figs<br />
653-656, <strong>10</strong>65, 1235, 1301) occurs on Sa/ix spp. and Popu/us tremu/a (Barnes, H.F.,<br />
1933); <strong>the</strong> host <strong>of</strong> <strong>the</strong> only o<strong>the</strong>r <strong>British</strong> species (P. errans Meigen) is unknown.<br />
The two <strong>British</strong> species <strong>of</strong> Ca/ycomyza are leaf (blotch) miners. C. humera/is (Roser)<br />
on Compositae (Asteraceae) especially Aster, Be/lis and Erigeron; C. artemesiae<br />
(Kaltenbach) on Artemisia and Eupa<strong>to</strong>rium.<br />
The genus Amauromyza (8 <strong>British</strong> species) contains leaf (blotch) miners and stemborers<br />
(s.g. Cepha/omyza).<br />
Of <strong>the</strong> 41 species <strong>of</strong> Liriomyza (figs 657-658) 34 are leaf miners; two (L. virgo<br />
(Zetterstedt), L.flavopicta Hendel) are stem-feeders, one feeds on flowerheads and <strong>the</strong><br />
hosts <strong>of</strong> 4 species are unknown. L. bryoniae (Kaltenbach) is probably an introduced<br />
species which forms an irregular linear mine (fig. 1299) and is a common pest <strong>of</strong> <strong>to</strong>ma<strong>to</strong><br />
(Lycopersicon escu/entum Miller) in glasshouses and also occurs on cucumbers<br />
( Cucumis sativus L.). L. pisivora Hering mines <strong>the</strong> leaves <strong>of</strong> peas (Pisum sativum L.) and<br />
L. congest a (Becker) mines <strong>the</strong> leaves <strong>of</strong> peas and beans ( Viciafaba L.) but <strong>the</strong>ir damage<br />
is negligible (Spencer 1973b ).<br />
Pteridomyza hi/are/la (Zetterstedt) is a leaf miner on <strong>the</strong> ferns Pteridium aqui/inum<br />
(L.) and Po/ypodium vulgare L.<br />
Of <strong>the</strong> four species <strong>of</strong> Me<strong>to</strong>pomyza three are known <strong>to</strong> be leaf-miners (two on Carex ,<br />
one on Viola). The hosts <strong>of</strong> M.flavonotata (Haliday) are probably Gramineae among<br />
which adults are not uncommon.<br />
The hosts <strong>of</strong> three <strong>of</strong> <strong>the</strong> 5 <strong>British</strong> species <strong>of</strong> Phy<strong>to</strong>liriomyza are known: P.<br />
me/ampyga (Loew) (formerly in Liriomyza), a blotch leaf-miner on Impatiens; P.<br />
arctica (Lundbeck), an external stem-miner on Sonchus and probably o<strong>the</strong>r<br />
Compositae; and P. pteridii Spencer ( 1973a) on Pteridium aqui/inum.<br />
No hosts are known for <strong>the</strong> three <strong>British</strong> Lemurimyza but <strong>the</strong> single <strong>British</strong><br />
Nemorimyza, N . posticata (Meigen), forms large blotch mines on Solidago.<br />
Of <strong>the</strong> 15 species <strong>of</strong> Paraphy<strong>to</strong>myza nine feed as leaf or external stem-miners<br />
on Rubiaceae (Ga/ium), Dipsacaceae (Knautia) and Caprifoliceae (Lonicera,<br />
Symphoricarpus) and six are leaf-miners on Salicaceae.<br />
Napomyza species are primarily internal stem-borers, pupating in <strong>the</strong> stem. The<br />
European N. carotae Spencer has not yet been recorded in Britain but <strong>the</strong> larvae cause<br />
damage <strong>to</strong> carrots superficially resembling that <strong>of</strong> <strong>the</strong> carrot fly (Psi/a rosae) and it<br />
could easily be overlooked (Spencer, 1973b ). N. lateralis (Fallen) (figs 659-661) is<br />
common in Compositae.<br />
Pseudonapomyza atra (Meigen) is a leaf miner on Gramineae (Avena, Lolium,<br />
Pha/aris, Poa, etc) and <strong>the</strong> only o<strong>the</strong>r <strong>British</strong> species, P. lacteipennis (Malloch) is<br />
certainly also a grass-feeder but as yet its hosts are unknown.<br />
112
Phy<strong>to</strong>myza (figs 664--667) is <strong>the</strong> largest genus <strong>of</strong> Agromyzidae in Britain. Eighty <strong>of</strong> its<br />
<strong>British</strong> species are leaf-miners, five feed mainly in <strong>the</strong> mid-rib, four are internal stem<br />
borers, six feed in seed heads and <strong>the</strong> biology <strong>of</strong> 11 species is unknown. Of <strong>the</strong> leafmining<br />
species P. syngenesiae (Hardy) and P. horticola Goureau are probably <strong>the</strong> best<br />
known. These two common species were formally confused under <strong>the</strong> name P. atricornis<br />
(<strong>of</strong> authors) until Griffiths (1967) revised this complex. Horticulturalists knew this<br />
name for <strong>the</strong> chrysan<strong>the</strong>mum leaf-miner (fig. 1303) which causes serious commercial<br />
losses <strong>to</strong> growers in Europe and <strong>the</strong> United States. This pest species is P. syngenesiae<br />
(figs 662-663, I 066, 1237) and in addition <strong>to</strong> chrysan<strong>the</strong>mums can cause serious<br />
damage <strong>to</strong> lettuces and is found on a wide range <strong>of</strong>Compositae and is perhaps mostly<br />
readily seen on Sonchus (though ano<strong>the</strong>r 8 species in 4 genera <strong>of</strong> agromyzids also occur<br />
on Sonchus!). P. horticola is more widely polyphagous than P. atricornis on Compositae<br />
and many o<strong>the</strong>r families and is an occasional pest on peas but in Britain has no<br />
serious effect on <strong>the</strong> crop. P. ilicis Curtis (<strong>the</strong> holly leaf-miner) (figs <strong>10</strong>67, 1238) is<br />
ano<strong>the</strong>r familiar species which produces blotch-mines on <strong>the</strong> leaves <strong>of</strong> flex aquifolium<br />
L.<br />
P. ru.fipes Meigen is a large species which feeds inside <strong>the</strong> stem or midrib <strong>of</strong> <strong>the</strong> larger<br />
leaves <strong>of</strong> Cruciferae, especially Brassica spp. on which it can be <strong>of</strong> some economic<br />
importance.<br />
Cerodontha is ano<strong>the</strong>r large genus, <strong>the</strong> larvae <strong>of</strong> which feed exclusively on<br />
monocotyledons. The hosts <strong>of</strong> <strong>the</strong> 34 <strong>British</strong> species are as follows: 12 on Gramineae,<br />
seven on Cyperaceae, two on Iridaceae, one on Juncaceae and 12 are unknown. C.<br />
( Dizygomyza) iridis (Hendel) is an abundant species which mines Iris foetidissima L.<br />
and ano<strong>the</strong>r common species, C. ( D.) ireos (Goureau), mines Iris pseudacorus L. (figs<br />
668, 1302).<br />
Chloropidae<br />
(Figs: larvae 669- 682, eggs <strong>10</strong>68-<strong>10</strong>69, puparia 1241-1242, galls 1305- 1306)<br />
Over 2,000 species <strong>of</strong> Chloropidae are known throughout <strong>the</strong> world but <strong>the</strong> family is<br />
poorly worked and many new species and genera undoubtedly await discovery and<br />
description and many life-his<strong>to</strong>ries require elucidation. The larvae <strong>of</strong> many species are<br />
phy<strong>to</strong>phagous, especially on Gramineae and several species are important pests <strong>of</strong><br />
cereals, such as frit flies (Oscinella) and gout flies (Chlorops). Some form galls and<br />
o<strong>the</strong>rs are saprophagous or even parasitic. Larvae <strong>of</strong> <strong>the</strong> Australian genus Batrachomyia<br />
live in perforated swellings under <strong>the</strong> skin <strong>of</strong> frogs. Some are preda<strong>to</strong>rs on root<br />
aphids, rice boring moth larvae, Thysanoptera, egg capsules <strong>of</strong> grasshoppers and egg<br />
sacs <strong>of</strong> spiders.<br />
Some 243 species in 39 genera and two subfamilies occur in Britain. Nye (1958) has<br />
described <strong>the</strong> larvae <strong>of</strong> several species living in Gramineae and provides a key, o<strong>the</strong>r<br />
papers on <strong>immature</strong> <strong>stages</strong> are cited below. Balachowsky & Mesnil (1935) is useful for<br />
chloropine larvae, especially <strong>the</strong> pest species. Viviparity has been shown <strong>to</strong> occur in<br />
some African Chloropidae (Spencer, 1985). General summaries <strong>of</strong> <strong>the</strong> biology <strong>of</strong> <strong>the</strong><br />
family in Europe are provided by Balachowsky & Mesnil (1935) and Wendt (1968).<br />
Valley, Wearsch & Foote (1969) discuss larval feeding habits <strong>of</strong> North American<br />
species.<br />
Oscinellinae. The three <strong>British</strong> species <strong>of</strong> Lipara cause galls (fig. 1305) in <strong>the</strong> common<br />
reed (Phragmites communis Trinius) and an account <strong>of</strong> <strong>the</strong>ir systematics, morphology,<br />
behaviour and ecology for all <strong>stages</strong> is given by Chvala et al. (1974). O<strong>the</strong>r dipterous<br />
larvae are found in Lipara galls, including Cecidomyiidae and o<strong>the</strong>r Chloropidae<br />
(noted below), some <strong>of</strong> which appear <strong>to</strong> be regular inquilines (Biair, 1932, 1944a,<br />
1944b).<br />
113
Collin ( 1946) records that Calamoncosis minima (Strobl) 'appears freely when<br />
breeding out Lipara lucens Meigen' and that C. glyceriae Nartshuk (as laminiformis<br />
(Becker)) feeds gregariously on <strong>the</strong> enshea<strong>the</strong>d inflorescence <strong>of</strong> <strong>the</strong> grass Glyceria<br />
aquatica (L.).<br />
Adults <strong>of</strong> Siphonella oscinina (Fallen) have been swept from conifers but <strong>the</strong><br />
<strong>immature</strong> <strong>stages</strong> are unknown. Coli in ( 1946) has reared Polyodaspis ruficornis<br />
(Macquart) from walnuts (probably imported). The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> two <strong>British</strong><br />
Goniopsita species appear <strong>to</strong> be unknown. Coli in ( 1946) has reared Lasiamba baliola<br />
(Collin) from 'material taken from an ulcerous elm tree'.<br />
Nye (1958) found <strong>the</strong> larvae <strong>of</strong> Conioscinella mimula Collin and C.frontella (Fallen)<br />
overwintering in <strong>the</strong> basal tillers <strong>of</strong> <strong>An</strong>thoxanthum odoratum L. and Holcus lanatus L.<br />
respectively. The adults <strong>of</strong> C. gallarum (Duda) appear <strong>to</strong> be associated with oak<br />
(Collin, 1946).<br />
Speccafrons halophila Duda has been reared from spiders' eggs.<br />
Gaurax niger (Czerny) has been reared from <strong>the</strong> nest <strong>of</strong> a dormouse by E. B. Basden<br />
(Collin, 1946). Larval and puparial characters <strong>of</strong> Gaurax dubius (Macquart) (figs<br />
669- 670) have been described (Smith, 1965) from puparia found in Poly porus betulinus<br />
Fries. G. fascipes Becker has been reared from nests <strong>of</strong> blackbird and linnet by<br />
E. B. Basden, but C. H. W. Pugh reared it from under bark in a dead branch and<br />
Smith (1967) casts doubt upon birds' nests as a usual habitat. Collin (1946) has<br />
reared lncertella zuercheri Duda from galls <strong>of</strong> both Lipara lucens and L. similis<br />
Schiner.<br />
Oscinella frit (L.) (figs 671-676, <strong>10</strong>68, 1241) (frit fly) is a serious pest <strong>of</strong> cereals,<br />
especially oats in Europe and mainly wheat in North America. It is a highly polyphagous<br />
species and is found in a wide variety <strong>of</strong>Gramineae. Steel (1931) and Nye<br />
(1958) describe <strong>the</strong> larvae <strong>of</strong> 0. frit, <strong>the</strong> latter including morphological variants in<br />
different hosts, and o<strong>the</strong>r closely related species in various Gramineae. Nartshuk's<br />
( 1956) paper on Oscine !la larvae should also be studied.<br />
Hapleginella laevifrons (Loew) has been reared from larch cones (D. Fergusson, in<br />
<strong>the</strong> <strong>British</strong> Museum (Nat. Hist.) collections).<br />
The larva <strong>of</strong> Elachiptera corn uta (Fallen) overwinters in <strong>the</strong> decaying leaves <strong>of</strong> Typha<br />
and is a follower <strong>of</strong> Oscinellafrit in infested cereal fields (Jepson & Southwood, 1960).<br />
Adults <strong>of</strong> E. megaspis (Loew) are found among watercress (Collin, 1946).<br />
Nothing appears <strong>to</strong> be known <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> following Oscinelline<br />
genera occurring in Britain: Aphanotrigonum, Dicraeus, Eribolus, Gampsocera,<br />
Lioscinella. Melanochaeta, Oscinomorpha. Oscinosoma, Siphunculina, Trachysiphonella.<br />
Tricimba. <strong>An</strong>yone rearing adults in <strong>the</strong>se genera would do well <strong>to</strong> establish at<br />
least <strong>the</strong> identity <strong>of</strong> <strong>the</strong> host plants if possible.<br />
Chloropinae. The agile larvae <strong>of</strong> Camarota curvipennis (Latreille) feed in <strong>the</strong> ears <strong>of</strong><br />
rye, wheat and barley (Balachowsky & Mesnil, 1935; Nye, 1958).<br />
Larvae <strong>of</strong> Platycephala planifrons (F.) develop in <strong>the</strong> stems <strong>of</strong> reed (Phragmites)<br />
(Wandolleck, 1899; Wendt, 1968).<br />
The large genus Meromyza contains at least 16 <strong>British</strong> species (seven were listed in<br />
<strong>the</strong> Check List, Smith 1976). The larvae <strong>of</strong> few species have been described (Fedoseeva,<br />
1966, gives a key, but in Russian) and few host plants are known but are probably<br />
included among those from which adults are commonly swept. With Ismay's (1981) key<br />
<strong>to</strong> adults a little careful field work should soon amend this situation. The larvae <strong>of</strong><br />
Meromyza femora/a Macquart and M. variegata Meigen (figs 677- 678) have been<br />
reared from Dactylis glomerata L. M. saltatrix (L.) is associated with a variety <strong>of</strong><br />
grasses and is a pest <strong>of</strong> wheat on <strong>the</strong> continent; M. pra<strong>to</strong>rum Meigen has been reared<br />
from marram grass (Ammophila).<br />
Cryp<strong>to</strong>nevraflavitarsis (Meigen) has been reared from Lipara galls (Biair, 1944b) but<br />
hosts <strong>of</strong> <strong>the</strong> o<strong>the</strong>r three species are unknown.<br />
114
The larvae <strong>of</strong> Lasiosina cinctipes (Meigen) is said <strong>to</strong> develop in barley following<br />
attacks by Chlorops pumilionis (Bjerkander).<br />
Larvae <strong>of</strong> Cetema neglecta Tonnoir overwinter in <strong>the</strong> basal tillers <strong>of</strong> Lolium perenne<br />
L. , Festuca pratensis Huds., Poa trivia/is L. and Agrostis spp. The similar C. elongata<br />
(Meigen) larva overwinters in Agrostis spp. and C. myopina Loew has been recorded<br />
from spring oats (probably from a ploughed-in ley) (Nye, 1958).<br />
According <strong>to</strong> Balachowsky & Mesnil (1935) <strong>the</strong> larva <strong>of</strong> Chlorops strigula (F.) overwinters<br />
in species <strong>of</strong> Brachypodium and Agropyron. Chlorops pumilionis (Bjerkander)<br />
( = taeniopus Meigen) (gout fly) (figs 679-682, <strong>10</strong>69, 1242) can be a serious pest <strong>of</strong><br />
cereals, particularly on <strong>the</strong> Continent. The larvae overwinter in <strong>the</strong> basal tillers <strong>of</strong><br />
wheat, barley, rye and Agropy ron re pens L. In this and o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> genus <strong>the</strong><br />
larvae stimulate <strong>the</strong> plant <strong>to</strong> hypertrophy and a swollen 'gouted' shoot is formed.<br />
Larvae <strong>of</strong> o<strong>the</strong>r species <strong>of</strong> Chlorops may be found in <strong>the</strong> following Gramineae: C.<br />
brevimana Loew in Phalaris arundinacea L.; C. interrupta Meigen in Agropyron spp.;<br />
and C. speciosa Meigen in Deschampsia caespi<strong>to</strong>sa L.<br />
The larva <strong>of</strong> Chloropisca glabra Meigen is predaceous on root-feeding aphids and <strong>the</strong><br />
same habit is shared by Thauma<strong>to</strong>myia not at a (Meigen), which feeds on <strong>the</strong> root aphid<br />
Pemphigus bursar ius (L.) and may be an important agent in biological control.<br />
No hosts appear <strong>to</strong> be known for <strong>the</strong> following chloropine genera occurring in<br />
Britain: Diplo<strong>to</strong>xa, Epichlorops, Eurina, Eutropha, Me/anum.<br />
Calyptratae<br />
Oestridae<br />
(Figs: larvae 683-692, puparium 1243)<br />
The family Oestridae contains 34 species in 9 genera and 2 sub-families found mainly<br />
in <strong>the</strong> Afrotropical and Palaearctic Regions. Four species in 3 genera representing both<br />
subfamilies occur in Britain.<br />
Larvae <strong>of</strong> Oestridae develop in <strong>the</strong> head cavities (nasal or pharyngeal) <strong>of</strong> species<br />
<strong>of</strong> mammals in several groups (Marsupialia, Proboscidea, Artiodactyla and<br />
Perissodactyla). The female flies deposit live first instar larvae (as many as 400--900)<br />
in<strong>to</strong> <strong>the</strong> nostrils <strong>of</strong> <strong>the</strong>ir hosts, which include several species <strong>of</strong> domestic animals, e.g.<br />
sheep, horse, goat and camel, and game animals. The three larval instars may differ<br />
markedly in appearance and as earlier instars are more likely <strong>to</strong> be involved in<br />
enquiries, <strong>the</strong>y are included in <strong>the</strong> illustrations (in this and <strong>the</strong> two following families).<br />
Zumpt (1965) gives a good general account <strong>of</strong> <strong>the</strong> life his<strong>to</strong>ries and morphology <strong>of</strong><br />
Oestridae and includes many references <strong>to</strong> o<strong>the</strong>r works. Papavero ( 1977) discusses <strong>the</strong><br />
classification <strong>of</strong> <strong>the</strong> family.<br />
Oestrinae. The first ins tar larvae (fig. 684) <strong>of</strong> Oestrus ovis L. (sheep nostril fly or sheep<br />
bot fly) are squirted in flight at <strong>the</strong> nostrils <strong>of</strong> sheep (also goats and dogs) where <strong>the</strong>y<br />
crawl in<strong>to</strong> <strong>the</strong> frontal sinuses. Here <strong>the</strong>y attach <strong>the</strong>mselves by <strong>the</strong>ir mouthhooks <strong>to</strong> <strong>the</strong><br />
mucous membrane and feed. They remain <strong>the</strong>re for some 9 months when <strong>the</strong> fully fed<br />
larvae (fig. 683) relinquish <strong>the</strong>ir hold on <strong>the</strong> membrane and are sneezed out by <strong>the</strong> sheep<br />
on<strong>to</strong> <strong>the</strong> ground where <strong>the</strong>y pupate (fig. 1243) under s<strong>to</strong>nes or tufts <strong>of</strong> grass. Usually<br />
only a few larvae are present in each animal, but <strong>the</strong>y may cause a serious loss <strong>of</strong><br />
condition. Occasionally large numbers may be found (up <strong>to</strong> 350 in one animal) and<br />
high mortality may occur (especially among lambs), usually during dry years.<br />
Occasionally (rarely in Britain) larvae may be deposited in <strong>the</strong> eye, mouth, nostrils<br />
or outer ear <strong>of</strong> man (especially people associated with sheep). Fortunately (unlike<br />
Hypoderma, see below), <strong>the</strong> larvae do not survive beyond <strong>the</strong> first instar but may cause<br />
115
what is usually diagnosed as acute catarrhal conjunctivitis (more correctly ophthalmomyiasis)<br />
for as long as <strong>10</strong> days. As many as 50 larvae have been removed from <strong>the</strong><br />
conjunctival sac <strong>of</strong> a single patient (Zumpt, 1965; Keiser, 1948). Although <strong>the</strong> larvae<br />
are very tiny, transparent, and easily overlooked, in <strong>the</strong>se cases <strong>the</strong> sclerotized mouthparts<br />
(fig. 686) are readily visible and identifiable even on rough slides or pho<strong>to</strong>graphs<br />
submitted for identification from hospitals. Cases occur in Britain but are rarely<br />
recorded in <strong>the</strong> literature and appear <strong>to</strong> occur in particular 'good' years (e.g. <strong>the</strong> dry<br />
summer <strong>of</strong> 1976 brought several cases).<br />
Cephenemyiinae. The main host <strong>of</strong> Cephenemyia auribarbis (Meigen) (<strong>the</strong> deer nostril<br />
fly) (figs 689- 690) is <strong>the</strong> red deer (Cervus e/aphus L.), and very occasionally <strong>the</strong> fallow<br />
deer (Dama dama (L.)) is infested. C. trompe (Modeer) (fig. 691) is found in <strong>the</strong> reindeer<br />
(Rangifer tarandus (L.)) and has been introduced with this host in<strong>to</strong> Scotland.<br />
Pharyngomyia picta (Meigen) (fig. 692) is a parasite <strong>of</strong> red deer and is regarded as<br />
extinct in Britain but as <strong>the</strong>re are said <strong>to</strong> be more deer in Britain now than for centuries<br />
it may well reappear (Smith, 1974a).<br />
Hypodermatidae<br />
(Figs: larvae 693-702, eggs <strong>10</strong>70--<strong>10</strong>72, puparium 1244)<br />
The family Hypodermatidae (warble-flies) contains 32 species in 11 genera and<br />
2 subfamilies, 29 <strong>of</strong> which occur in <strong>the</strong> Palaearctic Region. The family also occurs in<br />
<strong>the</strong> Afrotropical and Nearctic Regions. In <strong>the</strong> Check-list (Smith et al., 1976) <strong>the</strong><br />
Hypodermatidae is treated as a subfamily <strong>of</strong> <strong>the</strong> Oestridae but is now usually given<br />
family rank. One genus containing three species occurs in Britain.<br />
The larvae are specific parasites <strong>of</strong> mammals (Rodentia, Lagomorpha and<br />
Artiodactyla). The tiny eggs (figs <strong>10</strong>70--<strong>10</strong>72) (<strong>of</strong> which 300--800 may be laid) are glued<br />
<strong>to</strong> <strong>the</strong> hairs <strong>of</strong> <strong>the</strong> host. The first stage larvae penetrate <strong>the</strong> skin <strong>of</strong> <strong>the</strong> host usually<br />
by a hair follicle, and wander, ei<strong>the</strong>r through <strong>the</strong> connective tissue (subfamily Oestromyiinae<br />
- non-<strong>British</strong>) or internal organs (subfamily Hypodermatinae). The second<br />
and third stage larvae finally settle under <strong>the</strong> skin <strong>of</strong> <strong>the</strong> host and form cysts (warbles)<br />
with a hole <strong>to</strong> which <strong>the</strong>y apply <strong>the</strong>ir spiracles for respiration and through which <strong>the</strong><br />
mature larva eventually escapes. Larval feeding causes <strong>the</strong> host pain and discomfort<br />
which affect health and condition; <strong>the</strong> warbles damage <strong>the</strong> hide and thus give <strong>the</strong> family<br />
considerable economic importance when domestic animals are involved. Zumpt ( 1965)<br />
gives a good general account <strong>of</strong> <strong>the</strong> biology <strong>of</strong> warble flies .<br />
Hypoderma bovis (L.) (ox warble fly) (figs 693- 696) attacks cattle and occasionally<br />
horses. Eggs (figs <strong>10</strong>71-<strong>10</strong>72) are attached singly <strong>to</strong> <strong>the</strong> base <strong>of</strong> body hairs. Man is also<br />
sometimes attacked, both in <strong>the</strong> normal way when larvae are found in skin tumours and<br />
more seriously in <strong>the</strong> eye causing a malignant ophthalmomyiasis (see following<br />
species).<br />
Hypoderma lineatum (Villers) (lesser ox warble fly) (figs 697-700) also attacks cattle<br />
but in addition horses are occasionally infested. The eggs are laid in regular rows <strong>of</strong> 5 <strong>to</strong><br />
15 per hair (fig. I 070). Most <strong>of</strong> <strong>the</strong> records <strong>of</strong> warble-flies in man are probably <strong>of</strong> this<br />
species and it is possible that infestation occurs through handling cattle, ra<strong>the</strong>r than or<br />
as well as by direct oviposition. The few <strong>British</strong> cases known <strong>to</strong> me are recorded by<br />
Hope (1840), Style (1924), Smart (1939) and A. Smith & Greaves (1946). The most<br />
serious cases are those involving ophthalmomyiasis as (unlike Oestrus) <strong>the</strong> first stage<br />
larvae (fig. 698) may, if <strong>the</strong>y are not removed quickly, completely destroy <strong>the</strong> eyeball<br />
causing great pain (see Kriimmel & Brauns, 1956 and comments under Oestrus avis). I<br />
know <strong>of</strong> no <strong>British</strong> cases <strong>of</strong> ophthalmomyiasis involving Hypoderma. Again, infestation<br />
may occur through contamination (by wiping hands across eyes) after handling<br />
infested cattle ra<strong>the</strong>r than by direct oviposition, though <strong>the</strong> flies may possibly deposit<br />
116
eggs on eye-brows or lashes. The mouth parts <strong>of</strong> <strong>the</strong> first ins tar larvae are diagnostic and<br />
easily distinguishable from Oestrus by <strong>the</strong> <strong>to</strong>o<strong>the</strong>d mandibles (figs 696, 698).<br />
Hypoderma diana Brauer (deer warble-fly) (figs 70 1-702) develops in <strong>the</strong> roe deer but<br />
has also been found in fallow deer and has adapted <strong>to</strong> reindeer imported in<strong>to</strong> Scotland<br />
(Kettle & Utsi, 1955). Ironically, an initial period <strong>of</strong> quarantine had prevented <strong>the</strong><br />
<strong>introduction</strong> <strong>to</strong> Britain <strong>of</strong> <strong>the</strong> reindeer warble-fly (Oedemagena tarandi L.). As yet no<br />
records <strong>of</strong> human infestation by H. diana have been reported.<br />
Gasterophilidae<br />
(Figs: larvae 703- 711, eggs I 073-<strong>10</strong>76, puparium 1245)<br />
The family Gasterophilidae contains some 16-18 species in some 3-5 genera and two<br />
subfamilies. They are found in <strong>the</strong> Palaearctic, Afrotropical and Oriental regions but<br />
some Gasterophilus species have been accidentally introduced in<strong>to</strong> o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong><br />
world. Four species, all contained in <strong>the</strong> genus Gasterophilus, represent <strong>the</strong> family in<br />
Britain.<br />
The larvae are specific parasites <strong>of</strong> Perissodactyl mammals (horse and rhinoceros)<br />
and elephants (Proboscidea). Eggs (200- 2500) are deposited on skin and hairs (figs<br />
<strong>10</strong>73- <strong>10</strong>76) near <strong>the</strong> mouth, or in <strong>the</strong> grass. The larvae enter <strong>the</strong> mouth via <strong>the</strong> host's<br />
<strong>to</strong>ngue during body-licking, with <strong>the</strong> food, or by <strong>the</strong>ir own movement. Zumpt (1965)<br />
gives a good general account <strong>of</strong> <strong>the</strong> biology <strong>of</strong> <strong>the</strong> family and much <strong>of</strong> what follows is<br />
from his work, but based on non-<strong>British</strong> data.<br />
The normal hosts <strong>of</strong> Gasterophilus haemorrhoidalis (L.) (<strong>the</strong> nose bot fly) (fig. 703)<br />
are <strong>the</strong> horse and donkey (and zebra in Africa). The eggs (fig. <strong>10</strong>73) are brownish black<br />
with a stalk-like pedicel which is a continuation <strong>of</strong> <strong>the</strong> broad chorionic flange and<br />
hollow along one side <strong>to</strong> receive <strong>the</strong> supporting hair. The first instar larvae penetrate<br />
<strong>the</strong> lips and migrate in<strong>to</strong> <strong>the</strong> mouth. Second ins tar larvae are found in <strong>the</strong> s<strong>to</strong>mach and<br />
duodenum where <strong>the</strong>y moult <strong>to</strong> <strong>the</strong> third stage and eventually pass <strong>to</strong> <strong>the</strong> rectum where<br />
<strong>the</strong>y re-attach <strong>the</strong>mselves <strong>to</strong> <strong>the</strong> wall, <strong>of</strong>ten in great numbers, near <strong>the</strong> anus. Eventually<br />
<strong>the</strong>y detach <strong>the</strong>mselves and pass out, not necessarily with <strong>the</strong> faeces. Occasionally man<br />
may become infested with first instar larvae, usually on <strong>the</strong> face and but<strong>to</strong>cks, where<br />
<strong>the</strong>y cause a 'creeping myiasis' in <strong>the</strong> skin.<br />
G. intestinalis (De Geer) (<strong>the</strong> horse bot fly) (figs 705- 708) is only known from <strong>the</strong><br />
horse and donkey and <strong>the</strong>ir cross breeds. Occurrences in o<strong>the</strong>r animals (e.g. dogs,<br />
hyaenas, vultures) have only occurred when <strong>the</strong>y have fed on infested equine intestines.<br />
Eggs hatch on <strong>the</strong> application <strong>of</strong> moisture and friction and <strong>the</strong> first instar larvae travel<br />
on <strong>the</strong> <strong>to</strong>ngue and after some 24 days pass <strong>to</strong> <strong>the</strong> s<strong>to</strong>mach. Here <strong>the</strong>y complete <strong>the</strong>ir<br />
development <strong>to</strong> third instar, and are eventually excreted with <strong>the</strong> faeces and pupate in<br />
<strong>the</strong> soil. Records <strong>of</strong> 'creeping myiasis' in man may have involved o<strong>the</strong>r species <strong>of</strong><br />
Gasterophilus but <strong>the</strong>re is one probable North American record <strong>of</strong> human ophthalmomyiasis.<br />
G. nasalis (L.) (throat bot fly) (fig. 704) infests horses and donkeys (and Burchell's<br />
zebra). Eggs are attached <strong>to</strong> <strong>the</strong> hairs <strong>of</strong> <strong>the</strong> intermaxillary space between <strong>the</strong> rami <strong>of</strong><br />
<strong>the</strong> mandibles beneath <strong>the</strong> head. A female may lay some 300 <strong>to</strong> 500 eggs, usually one per<br />
hair (fig. <strong>10</strong>75), but occasionally up <strong>to</strong> 5 have been found. Undisturbed a female may<br />
lay 20 eggs on a given host. The larvae hatch without <strong>the</strong> stimulation <strong>of</strong> external<br />
moisture or pressure, migrate <strong>to</strong> <strong>the</strong> lips and invade <strong>the</strong> spaces between <strong>the</strong> teeth below<br />
<strong>the</strong> gum line and behind <strong>the</strong> alveolar process <strong>of</strong> <strong>the</strong> gums. Pus pockets form, following<br />
necrosis <strong>of</strong> <strong>the</strong> tissue, which may contain as many as 12 larvae. After 18-24 days <strong>the</strong><br />
larvae moult <strong>to</strong> <strong>the</strong> second instar and after a few more days move on <strong>to</strong> <strong>the</strong> duodenum.<br />
Here <strong>the</strong>y become attached <strong>to</strong> <strong>the</strong> wall near <strong>the</strong> pylorus and moult <strong>to</strong> <strong>the</strong> third stage.<br />
The larva takes some 11 months <strong>to</strong> mature and <strong>the</strong>n passes out with <strong>the</strong> faeces. Larvae<br />
are also said <strong>to</strong> occur in <strong>the</strong> s<strong>to</strong>mach. There are no records <strong>of</strong> 'creeping myiasis' in<br />
117
humans as <strong>the</strong> first ins tar larvae seem <strong>to</strong> be incapable <strong>of</strong> penetrating unbroken human<br />
skin.<br />
Normal hosts <strong>of</strong> G. pecorum (F.) (figs 709- 711) are <strong>the</strong> horse and donkey (and<br />
Burchell's zebra). The eggs are glossy black with a fringed pedicel and are not attached<br />
<strong>to</strong> <strong>the</strong> host's hairs but are laid on plants (fig. <strong>10</strong>76). A female lays from I ,300 <strong>to</strong> 2,500<br />
eggs which can remain alive for many months but once in <strong>the</strong> mouth <strong>of</strong> <strong>the</strong> host <strong>the</strong><br />
larvae hatch within 3- 5 minutes and immediately penetrate <strong>the</strong> mucous membrane <strong>of</strong><br />
<strong>the</strong> lips, gums, cheeks, <strong>to</strong>ngue or hard palate. They burrow <strong>to</strong>wards <strong>the</strong> root <strong>of</strong> <strong>the</strong><br />
<strong>to</strong>ngue and s<strong>of</strong>t palate, where <strong>the</strong>y remain for 9-<strong>10</strong> months until attaining <strong>the</strong> third<br />
ins tar when <strong>the</strong>y move <strong>to</strong> <strong>the</strong> s<strong>to</strong>mach. They may also be swallowed with <strong>the</strong> food and<br />
settle in <strong>the</strong> walls <strong>of</strong> <strong>the</strong> pharynx, oesophagus and s<strong>to</strong>mach. The fresh first stage larva<br />
(fig. 709) has a distinct crown <strong>of</strong> recurved spines on <strong>the</strong> first segment but <strong>the</strong>se disappear<br />
later (fig. 7<strong>10</strong>). First instar larvae can penetrate <strong>the</strong> skin within 3- 5 minutes and a<br />
human infestation can be easily obtained by passing <strong>the</strong> hand over grass on which flies<br />
have oviposited.<br />
A unique feature <strong>of</strong> <strong>the</strong> third stage larva is <strong>the</strong> arrangement <strong>of</strong> denticles on <strong>the</strong><br />
pseudocephalon in<strong>to</strong> 3 groups, 2 lying laterally and a third centrally in front <strong>of</strong> <strong>the</strong><br />
mouthhooks (fig. 711).<br />
This species has been much studied in Asia where it is regarded as <strong>the</strong> most<br />
pathogenic Gasterophilus species on horses (Zumpt, 1965).<br />
Tachinidae<br />
(Figs: larvae 712- 755, eggs <strong>10</strong>77- <strong>10</strong>85, puparia 1246-1251)<br />
This may be <strong>the</strong> largest family <strong>of</strong> Diptera although at present more species <strong>of</strong><br />
Tipulidae are actually known. Over 8000 species have been described. The classification<br />
<strong>of</strong> Tachinidae is fraught with difficulties because <strong>of</strong> <strong>the</strong> enormous number <strong>of</strong><br />
species and <strong>the</strong> variation in taxonomic value <strong>of</strong> <strong>the</strong> many morphological characters<br />
available on <strong>the</strong> adult flies. Herting (1984) recognises 1,552 species in 398 genera, 33<br />
tribes and 4 subfamilies for <strong>the</strong> Palaearctic Region. The <strong>British</strong> Check List (Smith et al. ,<br />
1976) listed some 236 species and <strong>the</strong> classification used here, for convenience, follows<br />
<strong>the</strong> sequence <strong>of</strong> genera <strong>of</strong> <strong>the</strong> Check List with higher taxonomic and nomencla<strong>to</strong>rial<br />
adjustments <strong>to</strong> conform with Herting's work, some faunistic additions, and <strong>to</strong> link with<br />
Emden's (1954) Handbook <strong>to</strong> adults.<br />
In spite <strong>of</strong> <strong>the</strong> varied form <strong>of</strong> <strong>the</strong> adults all Tachinidae share <strong>the</strong> parasi<strong>to</strong>id habit and<br />
most larvae live as endoparasites <strong>of</strong> o<strong>the</strong>r insects and rarely (Loewia and Eloceria)<br />
centipedes. The main hosts are Lepidoptera and Coleoptera, but Hemiptera,<br />
Hymenoptera, Orthoptera, Man<strong>to</strong>dea, Phasma<strong>to</strong>dea, and Embioptera are also<br />
attacked. Rarely Tachinidae are parasitic in <strong>the</strong>ir own order, usually in <strong>the</strong>ir larvae (e.g.<br />
Siphona), but only very rarely do <strong>the</strong>y attack adult flies (Smith, 1974c). However, as so<br />
few life-his<strong>to</strong>ries are known <strong>the</strong> possibility should always be borne in mind (see also<br />
Sarcophagidae, Mil<strong>to</strong>gramminae). Some tachinid groups are highly specialized in <strong>the</strong><br />
hosts <strong>the</strong>y select, while o<strong>the</strong>rs are polyphagous. Tachinidae may be oviparous or<br />
Iarviparous. Oviparous species may deposit large (macrotype) eggs (<strong>10</strong>0-200) on <strong>the</strong><br />
body <strong>of</strong> <strong>the</strong> host or small (microtype) eggs (2,000-6,000) on plants eaten by <strong>the</strong> host (see<br />
Salkeld, 1980). Larviparous species deposit first instar larvae on or near <strong>the</strong>ir hosts.<br />
Females <strong>of</strong> species that attack Hemiptera or adult Coleoptera may have <strong>the</strong>ir claspers<br />
and oviposi<strong>to</strong>rs specially adapted for piercing or specially modified spines for gripping<br />
<strong>the</strong> host. Some species no doubt exert a considerable control on certain agricultural<br />
pests and some have been deliberately used as biological control agents with varying<br />
success. In spite <strong>of</strong> this adequate descriptions <strong>of</strong> <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are few. Hennig<br />
(1948-1952) lists references <strong>to</strong> descriptions <strong>of</strong> under 300 species or about 3% <strong>of</strong> <strong>the</strong><br />
world fauna. For this reason, en<strong>to</strong>mologists should always preserve with full data any<br />
118
parasites reared from <strong>the</strong> insects <strong>the</strong>y are studying, including <strong>the</strong> puparial caps containing<br />
anterior spiracles and cast larval mouthparts (see Introduction) and pass <strong>the</strong>m<br />
on, with full host data, <strong>to</strong> interested specialists or suitable museums and institutions.<br />
The hosts <strong>of</strong> <strong>British</strong> Tachinidae are listed by Audcent ( 1942), Emden (1954) and<br />
Herting (1960). Papers by Hammond & Smith (1953- 1957) and Smith (1980) may be<br />
<strong>of</strong> interest <strong>to</strong> rearers <strong>of</strong> Lepidoptera and Emden (1950) <strong>to</strong> Coleopterists. For world<br />
species (as a host guide for unreared <strong>British</strong> species) see W. R . Thompson (1943-1965)<br />
continued by H erting ( 1971- ). Some useful papers describing <strong>immature</strong> Tachinidae are<br />
J. C. Nielsen (1909, 1911-1918), Bisset (1938), Zuska (1963), and Lehrer & Plugerj<br />
( 1966); o<strong>the</strong>r papers are mentioned below. Good accounts <strong>of</strong> <strong>the</strong> biology <strong>of</strong>Tachinidae<br />
are given by Clausen (1940) and Askew (1971). Ferrar (1987) should also be useful.<br />
Genera not mentioned may be assumed <strong>to</strong> be unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>. A new<br />
Handbook <strong>to</strong> <strong>the</strong> <strong>British</strong> Tachinidae is in active preparation by Robert Belshaw.<br />
Phasiinae<br />
Members <strong>of</strong> this subfamily are virtually exclusive parasites <strong>of</strong> Hemiptera<br />
Heteroptera. The egg (figs I 077- 1 079) is deposited on <strong>the</strong> host or injected through <strong>the</strong><br />
integument (Phasia). The mature larvae leave <strong>the</strong> host before pupation and <strong>the</strong> bug<br />
may survive for some days. The monograph <strong>of</strong> Dupius (1963) includes some <strong>immature</strong><br />
<strong>stages</strong>.<br />
Phasiini. Of <strong>the</strong> 3 <strong>British</strong> species <strong>of</strong> Phasia ( = A/ophora) <strong>the</strong> hosts <strong>of</strong> two are known:<br />
P. pusi//a Meigen which parasitizes Cydnus (Cydnidae) and Chi/acis (Lygaeidae), and<br />
P. obesa (F.) has been reared from Zicrona caeru/ea (L.) (Penta<strong>to</strong>midae). Cis<strong>to</strong>gaster<br />
species (figs 712- 714) are parasites <strong>of</strong> Penta<strong>to</strong>midae and some Cydnidae. Subclytia<br />
rotundiventris (Fallen) parasites Elasmucha grisea (L.) (Acanthosomatidae).<br />
Cylindromyiini. Cylindromyia brassicaria (F.) (figs 715- 716) parasitizes Penta<strong>to</strong>midae<br />
(Dolycoris, Holcostethus, Pa/omena), but <strong>the</strong> hosts <strong>of</strong> C. interrupta Meigen are<br />
unknown. Lophosia fasciata Meigen has been reared from Ae/ia (Penta<strong>to</strong>midae).<br />
Hemyda ( =£vibrissa) vittata (Meigen) has been reared from <strong>the</strong> penta<strong>to</strong>mid Arma<br />
cus<strong>to</strong>s Hahn (non-<strong>British</strong> species) on <strong>the</strong> Continent. Phania species have been reared<br />
from Carabidae (Col.) on <strong>the</strong> Continent.<br />
Leucos<strong>to</strong>matini. Cinochira atra Zetterstedt has been reared from Eremocoris p/ebeius<br />
(Fallen) (Lygaeidae). Foreign species <strong>of</strong> Dionaea and Leucos<strong>to</strong>ma parasitize Coreidae<br />
and <strong>the</strong> latter some o<strong>the</strong>r bugs as well.<br />
Herting (1984) includes <strong>the</strong> rare Redtenbacheria insignis Egger in <strong>the</strong> Phasiinae<br />
(Eu<strong>the</strong>rini) although its only recorded host is Lymantria monacha L.; it is also recorded<br />
from a pied flycatcher's nest! ·<br />
Dexiinae<br />
Dexiini. Members <strong>of</strong> <strong>the</strong> tribe Dexiini are virtually exclusive parasites <strong>of</strong> larval<br />
Coleoptera (<strong>the</strong> few records from Lepidoptera need confirmation and may be from<br />
species living in <strong>the</strong> same habitat as 'suitable' beetle hosts). The females deposit fully<br />
incubated eggs or first stage larvae in areas frequented by beetles, especially chafers.<br />
The larvae search for a host and probably gain entry through a spiracle. The mature<br />
larva leaves <strong>the</strong> host before <strong>the</strong> latter pupates.<br />
There are non-<strong>British</strong> records <strong>of</strong> Dinera carinifrons (Fallen) from larvae <strong>of</strong><br />
Lucanidae, Melolonthinae, Ce<strong>to</strong>niinae and Cerambycidae. D. grisescens (Fallen)<br />
parasitises Harpalus (Carabidae). Dexia species (Bovien & Bollwig, 1939) and Prosena<br />
siberita (F.) parasitise Melolonthinae.<br />
119
The two <strong>British</strong> species <strong>of</strong> Trixa are ovoviviparous and T. oestroidea (Robineau<br />
Desvoidy) has been recorded from Lepidoptera (Operophterafagata (Scharfenberg),<br />
Geometridae and Hepialus lupulinus (L.), Hepialidae). Only first instar larvae have<br />
been described (Zuska, 1962) and <strong>the</strong> genus clearly needs biological investigation.<br />
Herting (1984) includes <strong>the</strong> tribes Dufouriini and Voriini in Dexiinae (under his<br />
system Dufouriinae is <strong>the</strong> valid name for Dexiinae) which include genera placed in <strong>the</strong><br />
tribes Campylochaetini and Thelairini in <strong>the</strong> subfamilies Tachininae and Dufouriinae<br />
in <strong>the</strong> Check List.<br />
Dufouriini. The larvae <strong>of</strong> <strong>the</strong> tribe Dufouriini are mostly parasitic in adult Coleoptera,<br />
e.g. <strong>An</strong>thomyiopsis in Plagiodera (Chrysomelinae), Microsoma ( = Campogaster) (figs<br />
717-718) in Si<strong>to</strong>na (Curculionidae) (Muller, 1962), Dufouria (fig. 719) in Cassida<br />
(Chrysomelidae), Freraea in Carabidae, and Rondania, probably on weevils.<br />
Voriini. Members <strong>of</strong> this tribe are mostly parasites <strong>of</strong> Lepidoptera, with a few also<br />
attacking sawflies (Hym., Symphyta).<br />
The genera Voria, Athrycia and Cyr<strong>to</strong>phleba have been reared from Lepidoptera,<br />
mostly Noctuidae, and some species from <strong>the</strong> sawfly Diprion pini L.<br />
Blepharomyia pagana (Meigen) ( = amplicornis Zetterstedt) is parasitic on Geometridae<br />
(Bapta, Opisthograpta, Hybernia). Periscepsia carbonaria (Panzer), Ramonda<br />
( = Periscepsia in part) and Wagneria are parasitic on Noctuidae.<br />
Eriothrix species have only been reared on <strong>the</strong> Continent: E. prolixa (Meigen) from<br />
Pyrausta prophyralis Schilfermiiller (Pyralidae) and E. rufomaculata (De Geer) from<br />
Arctia hebe L. (Arctiidae), Noctuidae and 'Bombycids'.<br />
Campylochaeta praecox (Meigen) is a parasite <strong>of</strong> Crocallis elinguaria (L.) (Geometridae),<br />
several larvae developing in one caterpillar. C. ( = Elpe) inept a (Meigen)<br />
also parasitizes Geometridae, Sphingidae (Hyloicus pinastri (L.)), No<strong>to</strong>dontidae and<br />
(abroad) Arctiidae. Thelaira nigripes (F.) parasitizes caterpillars <strong>of</strong> larger moths,<br />
especially Arctiidae.<br />
Tachininae<br />
The tribal classification used here follows Herting (1984).<br />
Microphthalmini. Dexiosoma caninum (F.) is a parasite <strong>of</strong> cockchafer larvae<br />
(Melolontha, Coleoptera).<br />
Macquartiini. Macquartia species are parasites <strong>of</strong> Chrysomelidae (Col.) and are<br />
believed <strong>to</strong> deposit fully incubated eggs or newly hatched larvae in <strong>the</strong> vicinity <strong>of</strong> <strong>the</strong><br />
hosts. The parasite sometimes emerges only after pupation <strong>of</strong> <strong>the</strong> host.<br />
Pelatachinini. Pelatachina tibialis (Fallen) (figs 72G--721) is a parasite <strong>of</strong> Vanessa s.l.<br />
(Lepidoptera), also moths (Agrotidae, Aegeriidae).<br />
Ernestiini. The hosts <strong>of</strong> Zophomyia temula (Scopoli) are unknown. Hyalurgus lucidus<br />
(Meigen) is a sawfly parasite (Pteronidea, Pristiphora, Tenthredinidae). Gymnochaeta<br />
viridis (Fallen) parasitizes some Noctuidae, Geometridae and Lymantriidae,<br />
and Fausta nemorum (Meigen) has been reared from Orthosia cruda Denis &<br />
Schilfermiiller (Lep., Noctuidae). Ernestia species (fig. 722) parasitize caterpillars <strong>of</strong><br />
moths (Noctuidae, Lasiocampidae, etc.).<br />
Eurithia species parasitize Noctuidae and o<strong>the</strong>r moths; E. anthophila (Robineau<br />
Desvoidy) ( = radicum auctt.) has also been reared from <strong>the</strong> peacock butterfly (/nachis<br />
io (L.)) and various Sphingidae.<br />
Loewia foeda (Meigen) and Eloceria delecta (Meigen) are parasites <strong>of</strong> centipedes<br />
(Lithobius).<br />
120
Minthoini. Mintho ru.fiventris (Fallen) ( = lacera Rondani) is a parasite <strong>of</strong> Orthopygia<br />
glaucinalis L. (Pyralidae).<br />
Nemoraeini. Nemoraea pellucida (Meigen) is a parasite <strong>of</strong> Spilosoma lubricepida (L.)<br />
(Arctiidae), Bis<strong>to</strong>n betularia (L.) (Geometridae) and various Noctuidae.<br />
Leskiini. Aphria longirostris (Meigen) has been reared from 'cutworms' (Noctuidae<br />
larvae), and Bithia ( = Rhinotachina) modesta (Meigen) has been reared from an<br />
unnamed Sesiidae (Lep.); <strong>the</strong> host <strong>of</strong> B. spreta (Meigen) is unknown. Demoticus<br />
plebeius (Fallen) has been reared on <strong>the</strong> continent from Arctia hebe (L.) (Lep. Arctiidae),<br />
Solieria inanis (Fallen) from Spilosoma lutea Hufnagel (Arctiidae) and Orthosia<br />
incerta Hufnagel (Noctuidae); S. pacifica (Meigen) from <strong>the</strong> small <strong>to</strong>r<strong>to</strong>iseshell butterfly<br />
(Aglais urticae (L.)). Leskia aurea (Fallen) is a parasite <strong>of</strong> clearwing moths (Sesiidae)<br />
and is also recorded from <strong>the</strong> cod ling moth ( Cydia pomonella (L.), Tortricidae).<br />
Linnaemyini. Lypha dubia (Fallen) (figs 723- 725) is a parasite <strong>of</strong> <strong>the</strong> winter moth<br />
(Operophtera brumata (L.)) and Agriopis marginaria (F.), two important geometrid<br />
defolia<strong>to</strong>rs and o<strong>the</strong>r Lepidoptera. It has also been reared from <strong>the</strong> narcissus bulb fly<br />
(Merodon equestris, Syrphidae) (Collin, 1945). L. ru.ficauda (Zetterstedt) is a nor<strong>the</strong>rn<br />
species which has been reared from No<strong>to</strong>donta phoebe Siebert (No<strong>to</strong>dontidae) and<br />
Hydriomena impluviata Den is & Schiffermiiller ( = coerulata F.) (Geometridae).<br />
Lydina aenea (Meigen) has been reared from Panolis .fiammea (Denis & Schiffermiiller)<br />
( = griseovariegata Goeze) (Noctuidae) and Acrobasis tumidella (Zincken) (Pyralidae).<br />
Linnaemya species are also parasites <strong>of</strong> Lepidoptera, especially Noctuidae and<br />
Sphingidae, and Chrysosomopsis auratus (Fallen) attacks Horisme tersata (Denis &<br />
Schiffermiiller) (Geometridae).<br />
Tachinini. Peleteria rubescens Robineau-Desvoidy is a parasite <strong>of</strong> <strong>the</strong> swallowtail<br />
butterfly (Papilio machaon L.), various Noctuidae and <strong>the</strong> 'nun' moth, Lymantria<br />
monacha (L.) (Lymantriidae), which can be a serious defolia<strong>to</strong>r <strong>of</strong> trees on <strong>the</strong> Continent.<br />
Nowickiaferox (Panzer) is a parasite <strong>of</strong>Noctuidae. The large Tachina grossa (L.)<br />
is parasitic on Sphingidae, Lasiocampidae and Lymantriidae, and <strong>the</strong> o<strong>the</strong>r (smaller)<br />
species <strong>of</strong> Tachina (s.s.) use Lymantriidae and Noctuidae as hosts. T. (s.g. Servillia)<br />
lurida (F.) parasitizes Noctuidae and Sphingidae, but T. ( S.) ursina Meigen has yet <strong>to</strong><br />
be reared, as have any species <strong>of</strong> Germaria.<br />
Neaerini. On <strong>the</strong> continent Graphogaster brunnescens Villeneuve has been reared from<br />
Teleiodes notatella (Hiibner) (Gelechiidae), Evetria resinella (L.) (Tortricidae) and<br />
Leucoptera laburnella (Stain<strong>to</strong>n) (Lyonetiidae) (Herting, 1960). Phy<strong>to</strong>myptera nigrina<br />
(Meigen) ( =nitidiventris Rondani) parasitizes Geometridae, Pterophoridae and<br />
Olethreutidae. El.fia cingulata (Robineau-Desvoidy) is a parasite <strong>of</strong> Esperia sulphurella<br />
(F.) (Oecophoridae), <strong>the</strong> corn moth Nemapogon (=Tinea) granella (L.), and <strong>the</strong> cork<br />
moth N. ( = T.) cloacella (Ha worth) (Tineidae), and Neaera laticornis (Meigen) has<br />
been reared from Eucosmafulvana (Stephens) (Oiethreutidae).<br />
Triarthriini. Triarthria ( = Digonochaeta) setipennis (Fallen) ( = spinipennis Meigen)<br />
(figs 726-727) is a parasite <strong>of</strong> <strong>the</strong> common earwig For.ficula auricularia L. (Thompson,<br />
1928) (see also Ocytata, Goniini). Records from <strong>the</strong> caterpillars <strong>of</strong> various moths and<br />
beetles may be erroneous due <strong>to</strong> earwigs finding access <strong>to</strong> rearing cages or sharing larval<br />
tunnels. Fully incubated eggs are deposited near <strong>the</strong> hosts in<strong>to</strong> which <strong>the</strong> larvae<br />
burrow. The characteristic puparia (fig. 1249) may be found during winter in soil, under<br />
bark, etc.<br />
Siphonini. This tribe was included in <strong>the</strong> subfamily Goniinae in <strong>the</strong> Check List (Smith<br />
et al. 1976). First instar larvae <strong>of</strong> <strong>the</strong> tribe are described by O'Hara (1988).<br />
121
Actia species are parasitic on Tortricidae and o<strong>the</strong>r families <strong>of</strong> 'Microlepidoptera'<br />
including several pest species. A. lamia (Meigen) has been studied by Cheng (1967).<br />
Ceranthia, Ceromya, Goniocera and Peribaea species parasitize <strong>the</strong> larvae <strong>of</strong> larger<br />
moths e.g. Lasiocampidae.<br />
Siphona species parasitize Noctuidae (including Mamestra brassicae (L.)), Geometridae,<br />
etc., but S. cristata (F.) and S. geniculata (De Geer) have also been reared<br />
from lea<strong>the</strong>rjackets (Tipulidae larvae: T. maxima, T. oleracea and T. paludosa). S.<br />
geniculata (figs 728- 730) has 4 slits in each <strong>of</strong> <strong>the</strong> posterior spiracles. The larva is<br />
attached <strong>to</strong> one <strong>of</strong> <strong>the</strong> main tracheal trunks by means <strong>of</strong> a chitinous sheath (Rennie &<br />
Su<strong>the</strong>rland, 1920).<br />
Exoristinae<br />
Blondeliini. Admontia ( = Trichopareia) species are parasites <strong>of</strong> wood-boring lea<strong>the</strong>rjackets<br />
(Tipulidae larvae: Dictenidia, Ctenophora and Tanyptera). Belida angelicae<br />
(Meigen) has been reared from Tortrix viridana L. (Tortricidae), and Blondelia nigripes<br />
(Fallen) is parasitic on moths (Noctuidae, Geometridae, Lymantriidae, etc.) and<br />
occasionally on sawflies (Hymenoptera: Athalia, Pteronidea, Priophorus, Diprion).<br />
The common Compsilura concinnata (Meigen) (figs 731-732) has been reared from a<br />
greater variety <strong>of</strong> Lepidoptera than any o<strong>the</strong>r <strong>British</strong> tachinid; it also attacks sawfly<br />
larvae <strong>of</strong> several families. In <strong>the</strong> U .S.A. and Canada it has been introduced for use in<br />
<strong>the</strong> biological control <strong>of</strong> caterpillar defolia<strong>to</strong>rs including <strong>the</strong> nun moth (Lymantria<br />
monacha) and brown-tail moth (Euproctis chrysorrhoea (L.)) (Culver, 1919); it also<br />
attacks <strong>the</strong> cabbage white butterflies (Pieris brassicae (L.) and P. rapae (L.)) (Bissett,<br />
1938).<br />
The rare Leiophora innoxia (Meigen) is a parasite <strong>of</strong> adult fleabeetles (Haltica,<br />
Chrysomelidae), and Ligeria angusticornis (Loew) parasitizes plume moths<br />
(Pterophoridae). Medina species are parasites <strong>of</strong> adult Chrysomelidae (Coleoptera),<br />
but M. luctuosa (Meigen) has also been reared from some Lepidoptera.<br />
Meigenia species are parasites <strong>of</strong> Chrysomelid larvae, but M. mutabilis (Fallen)<br />
(figs 733- 736) has also been reared from sawfly larvae, moth larvae and Chorthippus<br />
(Orthoptera, Acrididae).<br />
Oswaldia muscaria (Fallen) attacks Geometridae and Noctuidae.<br />
Pericheta ( = Policheta) unicolor (Fallen) is recorded as a parasite <strong>of</strong> sawfly larvae<br />
(Croesus) and adult beetles (Chrysomela).<br />
Zaira cinerea (Fallen) (fig. 737) parasitizes adult Carabidae (Coleoptera).<br />
Exoristini. Bessa parasitizes many families <strong>of</strong> moths and numerous sawfly larvae.<br />
Diplostichus is also a sawfly parasite (Diprioniidae) and is recorded from Acronicta<br />
rumicis (L.) (Lep., Noctuidae).<br />
Exorista larvarum (L.) (figs 738-740) and E. grandis (Zetterstedt) are common<br />
parasites <strong>of</strong> a large number <strong>of</strong> families <strong>of</strong> butterflies and moths; E. fasciata (Fallen)<br />
occurs on Lymantriidae and Lasiocampidae but hosts are not known for <strong>the</strong> remaining<br />
<strong>British</strong> species <strong>of</strong> <strong>the</strong> genus.<br />
Parasetigena silvestris (Robineau-Desvoidy) is an important parasite <strong>of</strong> Lymantria<br />
monacha and L. dispar (L.) (<strong>the</strong> gipsy moth); it is also recorded from o<strong>the</strong>r Lymantriidae<br />
and Thyatiridae. Phorocera species are also recorded from gipsy moth and several<br />
o<strong>the</strong>r families <strong>of</strong> Lepidoptera. Chae<strong>to</strong>gena ( = S<strong>to</strong>ma<strong>to</strong>myia) acuminata (Rondani) is<br />
a moth parasite (Noctuidae) but also attacks terricolous Tenebrionidae (Blaps,<br />
Opatrum, etc., Coleoptera).<br />
Win<strong>the</strong>miini. Nemorillafloralis (Fallen) (figs 741-742) is a parasite <strong>of</strong>Tortricidae,<br />
Pyralidae, Oecophoridae (especially Depressaria), Hyponomeutidae, and many o<strong>the</strong>r<br />
families <strong>of</strong> moths, <strong>the</strong> peacock butterfly (Inachis io) and it has been recorded from<br />
122
a coccinellid beetle (Epilachna). Lehrer & Pascovici (1966) illustrate <strong>the</strong> <strong>immature</strong><br />
<strong>stages</strong>.<br />
Smidtia, Timavia and Win<strong>the</strong>mia parasitize several families <strong>of</strong> moths (e.g. Noctuidae,<br />
Geometridae, Lymantriidae, Lasiocampidae).<br />
Goniini. Genera in this tribe are mostly parasitic on Lepidoptera, with some on<br />
sawflies.<br />
Erycil/a ( = Al/ophorocera) ferruginea (Meigen) parasitizes Lymantria monacha and<br />
abroad is recorded from a noctuid (Hypena) and a <strong>to</strong>rtricid. The rare Brachichaeta<br />
strigata (Meigen) has been reared from No<strong>to</strong>donta dromedarius (L.) (No<strong>to</strong>dontidae).<br />
Clemelis pul/ata (Meigen) is a parasite <strong>of</strong>Pyralidae and abroad has been reared from a<br />
psychid.<br />
Cyzenis albicans (Fallen) is <strong>the</strong> most important parasite <strong>of</strong> <strong>the</strong> winter moth<br />
(Operophtera brumata). The female s<strong>to</strong>res as many as 2,000 minute black eggs in her<br />
greatly expanded uterus until <strong>the</strong>y are ready <strong>to</strong> hatch. The eggs are <strong>the</strong>n laid on leaves<br />
damaged by caterpillar feeding. Few <strong>of</strong> <strong>the</strong> eggs are in fact swallowed by winter moth<br />
caterpillars, but may be swallowed by <strong>the</strong> caterpillars <strong>of</strong> o<strong>the</strong>r species in which <strong>the</strong>y<br />
would not hatch, but most are not swallowed at all. The few successful eggs hatch in <strong>the</strong><br />
fore gut <strong>of</strong> <strong>the</strong> host and <strong>the</strong> larvae enter a cell <strong>of</strong> <strong>the</strong> salivary (silk) gland. O<strong>the</strong>r<br />
Geometridae are also attacked by Cyzenis.<br />
Elodia ambula<strong>to</strong>ria (Meigen) attacks Tineidae and Hyponomeutidae while E. morio<br />
(F alien) is a parasite <strong>of</strong> <strong>the</strong> codling moth ( Cydia pomonella (L. )), o<strong>the</strong>r Tortricidae and<br />
some o<strong>the</strong>r 'Microlepidoptera'.<br />
Erynnia ocypterata (Fallen) (omitted from <strong>the</strong> Check List) has been reared from<br />
Sparganothis pil/eriana (Denis & Schiffermiiller) (Tortricidae).<br />
Eumea linearicornis (Zetterstedt) ( = westermanni (Zetterstedt)) parasitizes Noctuidae,<br />
Pyralidae, Olethreutidae and Tortricidae.<br />
Eurysthaea scutellaris (Robineau-Desvoidy) (figs 743- 745) is one <strong>of</strong> <strong>the</strong> Tachinidae<br />
with 4 slits in <strong>the</strong> posterior spiracles (see also Siphona and key <strong>to</strong> families); it attacks<br />
Hyponomeutidae, Tortricidae, Noctuidae and Geometridae.<br />
The very rare Frontina laeta (Meigen) parasitizes Sphingidae and Philudoria<br />
pota<strong>to</strong>ria (L.) (Lasiocampidae); Masicera pavoniae (Robineau-Desvoidy) also attacks<br />
<strong>the</strong>se two families and is a parasite <strong>of</strong> <strong>the</strong> emperor moth (Saturnia pavonia (L.))<br />
(Saturniidae). Gonia species mostly parasitize Noctuidae. Hebia flavipes Robineau<br />
Desvoidy attacks Noctuidae and Geometridae, and Myxexoris<strong>to</strong>ps blondeli<br />
(Robineau-Desvoidy) parasitizes sawflies (Hym., Tenthrediniidae) and has also been<br />
recorded from Euproctis chrysorrhoea (L.) (Lymantriidae). Ocytata pal/ipes (Fallen)<br />
( = Rhacodineura antiqua (Meigen)) (figs 746-747) is a parasite <strong>of</strong> earwigs (Dermaptera,<br />
Forjicula- see also Triarthria) and is also recorded from <strong>the</strong> gipsy moth (Lymantria<br />
dispar) and Orthosia miniosa (Denis & Schiffermiiller) (Noctuidae).<br />
Pales pavida (Meigen) is commonly reared from Noctuidae, Lymantriidae and o<strong>the</strong>r<br />
families <strong>of</strong> Lepidoptera including butterflies (Jnachis io, Aglais urticae); it has also been<br />
reared from a sawfly, Tenthredo amoena Gravenhorst.<br />
Phryno vetula (Meigen) attacks Orthosia, Cosmia (Noctuidae) and Apocheima<br />
pilosaria (Denis & Schiffermiiller) (Geometridae) and Lymantria monacha. Platymya<br />
jimbriata (Meigen) parasitizes some Noctuidae, Geometridae, Zygaenidae and<br />
some sawflies. Thelymorpha marmorata (F.) attacks Lymantriidae, Lasiocampidae,<br />
Arctiidae, Noctuidae and has also been recorded from chrysomelid beetles<br />
(Phy <strong>to</strong>decta). Z enillia libatrix (Panzer) is ano<strong>the</strong>r wide-ranging parasite <strong>of</strong> many<br />
families <strong>of</strong> Lepidoptera.<br />
Eryciini. This tribe mostly parasitizes Lepidoptera (including several species <strong>of</strong><br />
butterflies) and some sawflies.<br />
123
Bactromyia aurulenta (Meigen) is a parasite <strong>of</strong> Strymonidia w-album (Knoch)<br />
(Lycaenidae) and moths <strong>of</strong> <strong>the</strong> families No<strong>to</strong>dontidae, Drepanidae, Noctuidae and<br />
Geometridae. Cadurciella tritaeniata (Rondani) parasitizes Callophrys rubi (L.)<br />
(Lycaenidae) and Ostrinia nubilalis (Hiibner) (Pyralidae). Carcelia species parasitize<br />
Lymantriidae, Lasiocampidae (including <strong>the</strong> lackey moth, Malacosoma neustria (L.)),<br />
Arctiidae, Noctuidae, etc. and some sawflies ( Cimbex, <strong>An</strong>antholyda). The large eggs <strong>of</strong><br />
Carcelia may be attached <strong>to</strong> <strong>the</strong> host by stalks (fig. <strong>10</strong>85).<br />
Senome<strong>to</strong>pia species (usually considered as a subgenus <strong>of</strong> Carcelia) are parasites <strong>of</strong><br />
Geometridae (including <strong>the</strong> pine-looper, Bupalus piniaria (L.) and Abraxas) and<br />
several o<strong>the</strong>r families <strong>of</strong> moths and (S. exisa (Fallen), figs 748-749) <strong>the</strong> brims<strong>to</strong>ne<br />
butterfly (Gonepteryx rhamni (L.), Pieridae) and Sphingidae.<br />
Thecocarcelia species are parasites <strong>of</strong> Hesperiidae and T. acutangulata (Macquart),<br />
recently added <strong>to</strong> <strong>the</strong> <strong>British</strong> List (Wyatt, N .P., 1986), has been reared from Thymelicus<br />
lineola (Ochsenheimer), <strong>the</strong> Essex skipper, in Austria (Car!, 1968). The very rare<br />
Drino Iota (Meigen) parasitizes certain Noctuidae (Acronicta, Xestia ashworthii<br />
(Doubleday), Orthosia stabilis, (Denis & Schiffermiiller)), Sphingidae, <strong>the</strong> lackey moth<br />
(Malacosma neustria, Lasiocampidae) and <strong>the</strong> large white butterfly (Pieris brassicae,<br />
Pieridae), whilst <strong>the</strong> very common Epicampocera succinct a (Meigen) (figs 75G--751)<br />
attacks <strong>the</strong> small white butterfly (Pieris rapae) and <strong>the</strong> emperor moth (Saturnia<br />
pavonia, Saturniidae). Eryciafuribunda (Zetterstedt) is a parasite <strong>of</strong> <strong>the</strong> marsh fritillary<br />
(Euphydryas aurinia (Rottemburg)) and Arctia caja (L.) (Arctiidae). Huebneria a/finis<br />
(Fallen) parasitizes many Arctiidae, Noctuidae, Lymantriidae, Lasiocampidae, <strong>the</strong><br />
emperor moth (Saturnia pavonia) and <strong>the</strong> butterflies Aglais urticae (Nymphalidae)<br />
and Callophrys rubi (Lycaenidae).<br />
The two very common species <strong>of</strong> Lydella are parasitic on a variety <strong>of</strong>Noctuidae and<br />
o<strong>the</strong>r families, including <strong>the</strong> economically important European corn-borer (Ostrinia<br />
nubilalis, Pyralidae). Nilea hortulana (Meigen) parasitizes Noctuidae, especially<br />
Acronicta species and several pest species <strong>of</strong> Lepidoptera including Mamestra brassicae<br />
(L.), Pieris brassicae, Malacosoma neustria, Orgyia antiqua (L.), etc.<br />
Phebellia species have been reared from Lymantriidae, Arctiidae and Noctuidae,<br />
and <strong>the</strong> sawflies Cimbex and Diprion. Probably most species occurring in Britain have<br />
been confused with P. glauca (Meigen) and <strong>the</strong>ir precise host ranges need clarification.<br />
Phryxe species (figs 752- 755) have been recorded from a wide range <strong>of</strong> Lepidopterous<br />
hosts <strong>of</strong> at least 15 families (as well as earwigs and sawflies) and are probably <strong>the</strong><br />
most commonly reared tachinids. However, <strong>the</strong>re has been confusion over <strong>the</strong> identity<br />
<strong>of</strong> <strong>the</strong> <strong>British</strong> species and no doubt records for P. vulgaris (Fallen) and P. nemea<br />
(Meigen) include some for o<strong>the</strong>r species.<br />
Pseudoperichaeta nigrolineata (Walker) ( = roseanae Brauer & Bergenstamm)<br />
parasitizes Pyralidae (including Ostrinia nubilalis, <strong>the</strong> European corn-borer),<br />
Tortricidae, Noctuidae and Lysandra bellargus (Rottemburg) (Lycaenidae). Tlephusa<br />
diligens (Zetterstedt) has been reared from Melanchra persicaria (L.) (Noctuidae), and<br />
Xylotachina diluta (Meigen) ( = ligniperdae Brauer & Bergenstamm) is a parasite <strong>of</strong> <strong>the</strong><br />
goat moth Cossus cossus (L.)<br />
Rhinophoridae<br />
(Figs: larvae 756-767, eggs <strong>10</strong>86-<strong>10</strong>87, puparium 1252)<br />
The family Rhinophoridae (woodlouse-flies) contains 23 species in l 0 genera mostly<br />
found in <strong>the</strong> Old World and best represented in <strong>the</strong> western Palaearctic Region and<br />
South Africa. Until recently <strong>the</strong> family has been variously placed as a subfamily <strong>of</strong><br />
ei<strong>the</strong>r Calliphoridae or Tachinidae. The larvae are parasitic in woodlice (Isopoda,<br />
Crustacea) and those <strong>of</strong> <strong>the</strong> first stage (figs 758, 764-767) are specially adapted for<br />
holding on<strong>to</strong> and penetrating <strong>the</strong> host (e.g. pseudopods, sclerotised scales, elongated<br />
124
pharyngeal sclerites, etc.). Eggs (figs <strong>10</strong>86--<strong>10</strong>87) are laid freely away from <strong>the</strong> host.<br />
Some species have been recorded from spiders (Arachnida) and bugs (Hemiptera) but<br />
<strong>the</strong>se associations require confirmation. Eleven species representing 8 genera occur in<br />
Britain.<br />
Crosskey (1977) provides a taxonomic review <strong>of</strong> <strong>the</strong> family. Thompson ( 1934) and<br />
Bedding ( 1973) have described <strong>the</strong> <strong>immature</strong> <strong>stages</strong> <strong>of</strong> seven <strong>of</strong> <strong>the</strong> <strong>British</strong> species: Phy<strong>to</strong><br />
melanocephala (Meigen) (figs 756--757), P. discrepans Pandellt!, Tricogena rubricosa<br />
(Meigen) (figs 758-760), Stevenia atramentaria (Meigen) (figs 766--767), Rhinophora<br />
lepida (Meigen), Paykullia maculata (Fallen) (figs 761 - 762) and Melanophora roralis<br />
(L.) (figs 763- 765).<br />
Bedding ( 1973) found that Porcellio scaber Latreille is much more heavily parasitized<br />
(by five species) than o<strong>the</strong>r equally common and gregarious woodlice, e.g. Oniscus<br />
eLSe/Ius L. (attacked by P. maculata and P. discrepans). Armadillidium vulgare (Latreille)<br />
is attacked by P. melanocephala and Trachelipus rathkei (Brandt) by S. atramentaria.<br />
No Rhinophoridae had been recorded from o<strong>the</strong>r <strong>British</strong> woodlice until Irwin (1985a)<br />
reared M . roralis from Porcellio spinicornis Say. Bedding found that among <strong>the</strong> species<br />
<strong>of</strong> Rhinophoridae attacking P. scaber specific habitats were favoured as follows:<br />
Melanophora on <strong>the</strong> upper shore, Phy<strong>to</strong> discrepans in rubbish dumps and gardens, and<br />
Paykullia on colonies <strong>of</strong> woodlice under loose bark.<br />
Fur<strong>the</strong>r research will clearly be fruitful on <strong>the</strong>se parasites <strong>of</strong> woodlice and <strong>the</strong><br />
excellent book by Sut<strong>to</strong>n (1972) will facilitate identification <strong>of</strong> <strong>the</strong> hosts. There is also<br />
an active <strong>British</strong> Isopoda Study Group organised from <strong>the</strong> Institute <strong>of</strong> Terrestrial<br />
Ecology, Huntingdon.<br />
Sarcophagidae<br />
(Figs: larvae 768- 785, eggs I 088-<strong>10</strong>90, puparia 1253-1254)<br />
The Sarcophagidae (flesh-flies) is a large and cosmopolitan family <strong>of</strong> about<br />
2,500 species in 5 subfamilies. Until recently <strong>the</strong> family was treated as a subfamily<br />
<strong>of</strong> <strong>the</strong> Calliphoridae. Some 56 species in 16 genera representing 4 subfamilies occur in<br />
Britain.<br />
The reproductive habit is mostly larviparous though some lay eggs. The larvae may<br />
be parasitic on Orthoptera, Isoptera, Hymenoptera and some Diptera, or saprophagous,<br />
and <strong>the</strong>y may (especially Wohlfahrtia) also be involved in cases <strong>of</strong> myiasis in<br />
man and animals, mostly abroad.<br />
W. R. Thompson ( 1920) describes <strong>the</strong> larvae <strong>of</strong> several species, especially first ins tar<br />
Mil<strong>to</strong>gramminae. Zumpt (1965) is useful for larvae <strong>of</strong> Sarcophaga. Some o<strong>the</strong>r<br />
references are cited below and <strong>the</strong> excellent work <strong>of</strong> Pape (1987) should be consulted.<br />
Mil<strong>to</strong>gramminae. Larvae <strong>of</strong> Amobia signata (Meigen) (figs 768-770) are found in <strong>the</strong><br />
nests <strong>of</strong> wasps (Sphecidae, Vespidae) and some bees (Apidae). The adult female settles<br />
near <strong>the</strong> nest and enters as <strong>the</strong> host leaves. Inside <strong>the</strong> nest she deposits small larvae<br />
which commence feeding on <strong>the</strong> food s<strong>to</strong>re <strong>of</strong> <strong>the</strong> host larvae (e.g. spiders, caterpillars,<br />
etc.).<br />
Adults <strong>of</strong> Mil<strong>to</strong>gramma species (figs 771) shadow <strong>the</strong> host sand or digger wasp<br />
(Sphecidae) back <strong>to</strong> <strong>the</strong> nest, <strong>the</strong>n dart in <strong>to</strong> lay one or two eggs on <strong>the</strong> prey, <strong>the</strong>n<br />
quickly exit.<br />
Larvae <strong>of</strong> Senotainia conica (Fallen) (figs 772- 774) also develop in <strong>the</strong> nests <strong>of</strong><br />
Sphecidae. Adult females apparently oviposit on <strong>the</strong> female wasps while <strong>the</strong>y are<br />
carrying prey back <strong>to</strong> <strong>the</strong> nests.<br />
Larvae <strong>of</strong> Pterella grisea (Meigen) have been found in Cerceris (Hym., Sphecidae)<br />
burrows in Clythra (Col., Chrysomelidae) collected by <strong>the</strong> female wasp as food for its<br />
larvae.<br />
125
Me<strong>to</strong>pia species (fig. 775) are apparently viviparous. The female ei<strong>the</strong>r enters <strong>the</strong> nest<br />
( <strong>of</strong>Sphecidae, Apidae and Pompilus) or pounces upon <strong>the</strong> prey as it is being dragged in.<br />
The larvae <strong>of</strong> Oebalia ( = Ptychoneura) (figs 776-777) are found in <strong>the</strong> nests <strong>of</strong><br />
Sphecidae (Coelocrabro, Cemonus, Rhopalum, Pemphredon). Eggs are laid on <strong>the</strong><br />
female wasps (figs I 088-1 089) (Day & Smith, 1980) and this behaviour has also been<br />
observed in North America (Sanborne, 1982).<br />
Macronychiinae. Macronychia striginervis (Zetterstedt) ( = ungulans (Pandelle))<br />
larvae have been found in <strong>the</strong> nests <strong>of</strong> Vespidae, Sphecidae and Bombus. In North<br />
America P. H. Thompson (1978) has found larvae <strong>of</strong> a Macronychia species (near<br />
aurata Coquillett) in adult Tabanidae. Dipterous parasites <strong>of</strong> Diptera are very rare<br />
indeed, especially <strong>of</strong> adults (see under Tachinidae and Smith, 1974c).<br />
The female <strong>of</strong> Brachicoma de via (Fallen) deposits young larvae in Bombus (especially<br />
B. pascuorum (Scopoli) ( =agrorum (F.)) and Vespula nests, where <strong>the</strong>y are very<br />
common. The larvae (figs 778- 779) attack and kill <strong>the</strong> prepupal stage <strong>of</strong> <strong>the</strong> host and<br />
pupate in <strong>the</strong> nest material (summer pupae) or in nearby soil (hibernating pupae).<br />
Agriinae. Nyctia ha/terata (Panzer) is parasitic in weevils (Lixus). Larvae <strong>of</strong><br />
<strong>An</strong>giome<strong>to</strong>pa falleni Pape (=ruralis Fallen) are said <strong>to</strong> have been found abroad in<br />
superficial wounds <strong>of</strong> man and horse. The larvae <strong>of</strong> Agria species are polyphagous<br />
and frequently parasitic or predaceous on insects, especially <strong>the</strong> larvae and pupae <strong>of</strong><br />
Lepidoptera (Tolg, 1913). Sarcophila latifrons (Fallen) (fig. 780) feeds on carrion,<br />
dead insects and possibly also living insects. It has been recorded from various<br />
Acrididae.<br />
Helicobosca distinguenda Villeneuve is viviparous, producing larvae some 5 mm long<br />
which feed on dead snails. Some workers regard this genus as more correctly placed in<br />
<strong>the</strong> Calliphoridae.<br />
Sarcophaginae. Ravinia pernix (Harris) is parasitic in molluscs and insects.<br />
Blaesox ipha species are ovoviviparous; <strong>the</strong> larvae are parasitic in grasshoppers and<br />
locusts (Orthoptera, Acrididae) abroad (see Grea<strong>the</strong>ad, 1963 who reviews <strong>the</strong> parasites<br />
<strong>of</strong> Acrididae).<br />
Sarcophaga species (figs 781- 785) are larviparous and have a wide range <strong>of</strong> larval<br />
habitats from saprophagous feeding in decaying organic matter, dung and carrion <strong>to</strong><br />
parasitism (<strong>of</strong> molluscs, insects and spider egg sacs) and causing myiasis in man and<br />
o<strong>the</strong>r vertebrates, as follows:<br />
Dung (including human excrement): S. a/biceps Meigen, S. argyros<strong>to</strong>ma Robineau-Desvoidy,<br />
S. cruentata Meigen (=haemorrhoidalis (Fallen)) (figs 781- 785), S. exuberans Pandelle, S.<br />
incisi/obata Pandelle.<br />
Carrion: S. argyros<strong>to</strong>ma, S. carnaria L., S. cruentata.<br />
Myiasis: S. argyros<strong>to</strong>ma, S. carnaria, S. exuberans, S. cruentata.<br />
Molluscs (Helix, etc.): S. agnata Rondani, s: anaces Walker ( = setipennis Rondani), S.<br />
argyros<strong>to</strong>ma, S. filia Rondani, S. haemorrhoa Meigen, S. hirticrus Pandelle, S. me/anura<br />
Meigen, S. nigriventris Meigen (from living He/ice/la and Theba) and S. teretirostris Pandelle.<br />
Insects: S. a/biceps (Lepidoptera; Scarabaeidae and Saperda (Col.)), S. aratrix Pandelle<br />
(Lymantria monacha (Lep.), Prionus (Col.), S. argyros<strong>to</strong>ma (locusts)), S. caerulescens<br />
Zetterstedt ( =scoparia Pandelle) (Lymantria monacha), S. exuberans (Lepidoptera, Diprion<br />
(Hym.), Orthoptera), S. incisilobata (locusts), S. nigriventris ('locust'; Carabus, moribund<br />
Nicrophorus, Blaps (Col.)).<br />
Spider egg sacs (Epeira cornuta Koch): S. sexpunctata F. ( = clathrata Meigen).<br />
The above data cover <strong>the</strong> whole zoogeographical range <strong>of</strong> <strong>the</strong> species concerned and<br />
include non-<strong>British</strong> records (especially <strong>of</strong> myiasis). However, some identifications may<br />
be suspect; for example <strong>the</strong> common S. carnaria has been widely reported <strong>to</strong> cause<br />
126
myiaSIS m man (e.g. James, 1947). Zumpt (1965) refutes this, citing Kirchberg's<br />
claim (1954) that S. carnaria is an obligate parasite <strong>of</strong> earthworms. Greenberg<br />
( 1971: 79) however casts doubt on Kirchberg's claim as he has reared S. carnaria from<br />
carnon.<br />
Clearly, detailed study <strong>of</strong> <strong>the</strong> precise feeding habits <strong>of</strong> <strong>the</strong> larvae <strong>of</strong> our <strong>British</strong><br />
Sarcophaga would be a useful undertaking providing extreme care were taken over<br />
identification (from reared adult males which should be submitted <strong>to</strong> a specialist).<br />
Calliphoridae<br />
(Figs: larvae 786--790, 794-828; eggs 791, <strong>10</strong>91-<strong>10</strong>93; puparia 792-793, 1255-1256)<br />
Over 1<strong>10</strong>0 species <strong>of</strong> Calliphoridae (blow-flies) are known and <strong>the</strong> family occurs<br />
abundantly in all zoogeographical regions. The classification is difficult and <strong>the</strong> family<br />
is variously divided in<strong>to</strong> 5 <strong>to</strong> I 0 subfamilies by different workers. Some 32 species in 11<br />
genera representing 4 subfamilies occur in Britain.<br />
Calliphoridae are oviparous or larviparous. The larvae (maggots) may be<br />
omnivorous, carnivorous or parasitic on or in decaying organic matter such as carrion,<br />
excrement, foodstuffs, insects, earthworms, snails, <strong>to</strong>ads, birds and mammals. They<br />
may also cause myiasis in man and o<strong>the</strong>r vertebrates. The nomenclature follows<br />
Schumann (in So6s & Papp, 1986) but for convenience <strong>the</strong> sequence <strong>of</strong> genera follows<br />
<strong>the</strong> <strong>British</strong> Check List (Smith 1976).<br />
Calliphorinae. Larvae <strong>of</strong> Calliphora (bluebottles) occur in a variety <strong>of</strong> decaying<br />
substances, especially carrion, including human cadavers which gives <strong>the</strong>m considerable<br />
forensic importance in establishing postmortem interval (time <strong>of</strong> death) (Smith<br />
1986b ). Their occurrence in human foods, especially meat products, cheese, etc. makes<br />
<strong>the</strong>m easily <strong>the</strong> commonest insect larvae submitted <strong>to</strong> en<strong>to</strong>mologists for identification.<br />
Usually accompanying such requests is a desire <strong>to</strong> know rate <strong>of</strong> development and<br />
temperature-<strong>to</strong>lerance (especially under refrigeration). A few comments on this may<br />
<strong>the</strong>refore be <strong>of</strong> value. Calliphora oviposits at night (except in slaughterhouses) and it is<br />
doubtful if eggs are laid at temperatures lower than l2°C (though oviposition on partly<br />
frozen pig carcasses when exposed <strong>to</strong> air temperatures has been observed). At temperatures<br />
below 4°C Calliphora eggs will not hatch but at 6--rC hatching <strong>of</strong> eggs and larval<br />
development occurs. Up <strong>to</strong> 300 eggs may be laid, ei<strong>the</strong>r in smaller groups or a single<br />
batch, in natural orifices, crevices in <strong>the</strong> surface or wounds. The rate <strong>of</strong> development<br />
fluctuates with temperature but at 27°C and 50% R. H. for C. vicina Robineau<br />
Desvoidy (<strong>the</strong> common urban bluebottle) is recorded as follows (average in brackets):<br />
egg, 20-28 (24) hours; first stage larva, 18-34 (24) hours; second stage larva 16--28 (20)<br />
hours; third stage larva, 30-68 (48) hours; prepupa, 72- 290 (128) hours; pupa, 9- 15<br />
(!I) days; <strong>to</strong>tal <strong>immature</strong> <strong>stages</strong> 14-25 (18) days (figures after Kamal, 1958). On<br />
completion <strong>of</strong> feeding <strong>the</strong> maggots disperse from <strong>the</strong> food source up <strong>to</strong> a distance <strong>of</strong> 20<br />
feet or more searching for pupation sites. When this occurs indoors, this may result in<br />
maggots being sent <strong>to</strong> en<strong>to</strong>mologists via local health authorities. In such cases hidden<br />
sources should be sought, e.g. dead pigeons trapped behind sealed fireplaces, dead mice<br />
in attics, cellars, cupboards, etc. To facilitate identification <strong>of</strong> this very important<br />
species all <strong>the</strong> <strong>immature</strong> <strong>stages</strong> are shown (figs 786--793) on one plate and some also <strong>to</strong><br />
illustrate <strong>the</strong> introduc<strong>to</strong>ry sections.<br />
C. vomi<strong>to</strong>ria (L.) (<strong>the</strong> rural bluebottle) may be equally common, especially in<br />
country areas, and has a similar life-his<strong>to</strong>ry, but appears <strong>to</strong> spend longer in each <strong>of</strong> <strong>the</strong><br />
later <strong>stages</strong> (especially <strong>the</strong> prepupa) (see Kamal, 1958); fur<strong>the</strong>r investigation is<br />
required.<br />
O<strong>the</strong>r Calliphora species share similar larval habits but are more restricted in<br />
distribution. Calliphora larvae may be distinguished at once from Lucilia species<br />
127
(except L. ampullacea Villeneuve, fig. 81 0) found in similar situations by <strong>the</strong> presence <strong>of</strong><br />
<strong>the</strong> accessory oral sclerite (os) between <strong>the</strong> mandibles (best seen in ventral view).<br />
Erzin
Phormiinae. Phormia regina (Meigen) and Pro<strong>to</strong>phormia terranovae (Robineau<br />
Desvoidy) (figs 819- 822) are both carrion feeders and myiasis causers. Although P.<br />
terranovae has not been recorded as causing myiasis in man, both species have been<br />
used <strong>to</strong> clean wounds sustained by soldiers (see comments under Lucilia sericata). Both<br />
species may have forensic importance in human cadavers (Smith, 1986b).<br />
The larvae <strong>of</strong> Pro<strong>to</strong>calliphora azurea Fallen (bird blowfly) (figs 823- 824) occur in <strong>the</strong><br />
nests <strong>of</strong> birds (swallow, sparrows and o<strong>the</strong>r birds especially Oscine Passeriformes)<br />
where <strong>the</strong>y suck <strong>the</strong> blood <strong>of</strong> nestlings by means <strong>of</strong> a suction pad on <strong>the</strong> first segment,<br />
<strong>of</strong>ten killing <strong>the</strong>m.<br />
Rhiniinae. S<strong>to</strong>morhina lunata (F.) (figs 825-828) has only been recorded sporadically<br />
in Britain (1901 , with scattered records <strong>to</strong> 1947). Abroad it lays its eggs in <strong>the</strong> egg pods<br />
<strong>of</strong>locusts (Grea<strong>the</strong>ad, 1963) and appears <strong>to</strong> be associated with locust swarms.lt may be<br />
significant that in <strong>the</strong> two years cited <strong>the</strong>re was notable insect immigration in this<br />
country, 1947 in particular yielding many records <strong>of</strong> <strong>the</strong> migra<strong>to</strong>ry locust (Locusta<br />
migra<strong>to</strong>ria (L.)) (Colyer & Hammond, 1951). Perhaps it may adapt (or has adapted) <strong>to</strong><br />
a <strong>British</strong> Acridid.<br />
Scathophagidae<br />
(Figs: larvae 829- 854, eggs I 094-<strong>10</strong>96, puparia 1257-1262, leaf-mines 1307- 1308)<br />
Some 360 species <strong>of</strong> Scathophagidae in 66 genera are distributed mainly in <strong>the</strong><br />
Nor<strong>the</strong>rn Hemisphere. The larvae are mostly phy<strong>to</strong>phagous but some develop in dung,<br />
possibly in carrion, and o<strong>the</strong>rs in rotting seaweed on <strong>the</strong> coast. Fifty-three species in 22<br />
genera and 2 subfamilies occur in Britain. <strong>British</strong> genera not mentioned may be<br />
assumed <strong>to</strong> be unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>.<br />
Scathophaginae. Norellia spinipes (Meigen) (figs 829- 832) has been found <strong>to</strong> mine <strong>the</strong><br />
leaves <strong>of</strong> daffodils (Narcissus) (fig. 1308), pupating at <strong>the</strong> base <strong>of</strong> <strong>the</strong> plant and sometimes<br />
damaging <strong>the</strong> bulbs (Ciampolini, 1957, Chandler & Stubbs, 1969, 1970; de Jong,<br />
1985; Smith & Vardy, 1988). N. (s.g. Norellisoma) spinimana (Fallen) (figs 833-834)<br />
mines stems <strong>of</strong> docks (Rumex) (Disney, 1976b) and N. (N.) liturata (Meigen) appears <strong>to</strong><br />
be associated with meadowsweet (Filipendula).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Cordilura species develop in Carex, Scirpus and ]uncus in <strong>the</strong><br />
U.S.A. according <strong>to</strong> Wallace & Neff (1971) who include one Holarctic species that<br />
occurs in Britain, viz. C. pudica Meigen (figs 835- 837). The larvae <strong>of</strong> Cordilura are<br />
apparently distinguished by <strong>the</strong> presence <strong>of</strong> paired bifid mouth-hooks (fig 838) which<br />
should help in searching for <strong>the</strong> <strong>10</strong> o<strong>the</strong>r <strong>British</strong> species so far unknown in this stage.<br />
Nanna ( = Amaurosoma) species (timothy flies) (figs 839-840) feed on <strong>the</strong> flower<br />
heads <strong>of</strong> grasses and some (N. armillata (Zetterstedt), N.jlavipes (Fallen)) are pests <strong>of</strong><br />
rye and Phleum pratense L. (Timothy grass) (Nye, 1958).<br />
The larvae <strong>of</strong> Cleigastra ( = Cnemopogon) apicalis (Meigen) occur in Phragmites<br />
where <strong>the</strong>y were thought <strong>to</strong> be predaceous on <strong>the</strong> larvae <strong>of</strong> Lipara (Chloropidae) and<br />
where <strong>the</strong>ir puparia overwinter between <strong>the</strong> <strong>to</strong>p blades <strong>of</strong> <strong>the</strong> Lipara galls (Chvala et al.,<br />
1974). However, Groth ( 1969) found that <strong>the</strong> Cleigastra larvae feed on <strong>the</strong> excrement <strong>of</strong><br />
caterpillars (Archanara geminipuncta Haworth and Arenos<strong>to</strong>la phragmitidis Hiibner,<br />
Noctuidae) in <strong>the</strong> stems.<br />
According <strong>to</strong> Vockeroth (in Chandler & Stubbs, 1974), Cosme<strong>to</strong>pus, Microprosopa,<br />
Pogonota, Chae<strong>to</strong>sa, Trichopalpus and Spaziphora belong <strong>to</strong> a group <strong>of</strong> genera which<br />
probably have predaceous larvae living in subaquatic or aquatic situations. Of this<br />
group, only <strong>the</strong> life cycle <strong>of</strong> Spaziphora ( = Spathiphora) hydromyzina (Fallen) (figs<br />
841, 842) had <strong>the</strong>n been fully worked out. It occurs in sewage beds and stagnant water<br />
where <strong>the</strong> larva grazes over <strong>the</strong> pebbles, on algae and fungal growth but also eats<br />
129
cocoons <strong>of</strong> Chironomidae and clusters <strong>of</strong> Lumbricillus lineatus (Muller) worms and<br />
cocoons (Graham, 1939; Lloyd et al., 1940).<br />
The genus Acanthocnema can now be added <strong>to</strong> this group <strong>of</strong> aquatic Scathophagidae.<br />
Females <strong>of</strong> Acanthocnema ( Clinoceroides) glaucescens (Loew) crawl beneath <strong>the</strong> water<br />
surface <strong>of</strong> streams, down <strong>the</strong> sides <strong>of</strong> projecting s<strong>to</strong>nes. On <strong>the</strong> underside <strong>of</strong> <strong>the</strong> s<strong>to</strong>nes<br />
<strong>the</strong>y lay <strong>the</strong>ir eggs in <strong>the</strong> egg masses <strong>of</strong> Dixa, o<strong>the</strong>r flies and Trichoptera. The larva feeds<br />
on <strong>the</strong> eggs and surrounding jelly <strong>of</strong> <strong>the</strong> host (Hin<strong>to</strong>n, 1981 ). Coniosternum obscurum<br />
(Fallen) (formerly in Scathophaga) may now also be included in this group. The larvae<br />
feed on <strong>the</strong> egg masses <strong>of</strong> caddis flies (Trichoptera) (Berte & Wallace, 1987).<br />
Scathophaga stercoraria (L.) (<strong>the</strong> yellow dung fly) (figs 843- 84 7) breeds in dung (cow,<br />
sheep, horse, dog, poultry and human) where <strong>the</strong> larvae are said <strong>to</strong> be carnivorous.<br />
The larva is described by Schumann (1960). O<strong>the</strong>r dung breeding Scathophaga<br />
include S. furcata (Say) (dog, human, sheep, privies; also owl pellets); S. lutaria (F.)<br />
(human); S. inquinata Meigen and S. scybalaria (L.) (unspecified 'dung'). S. li<strong>to</strong>rea<br />
Fallen, S. calida Curtis and Ceratinos<strong>to</strong>ma ostiorum (Curtis) breed in thick moist rotten<br />
seaweed on <strong>the</strong> sea shore.<br />
Gimnomera tar sea (Fallen) (fig. 848) develops in <strong>the</strong> seed pods <strong>of</strong> Pedicularis species<br />
(Scrophulariaceae) (Chandler, 1975).<br />
The larvae <strong>of</strong> Hydromyza livens (F.) (figs 849-851) mine <strong>the</strong> stems and leaves <strong>of</strong> <strong>the</strong><br />
yellow water lilies (Nymphaea, Nuphar) (fig. 1307).<br />
Delininae. Parallelomma vittatum (Meigen) and Delina nigrita (Fallen) (figs 852-854)<br />
are both leaf-miners in orchids (Meijere, 1940). The life-his<strong>to</strong>ry <strong>of</strong> Lep<strong>to</strong>pa ftliformis<br />
Zetterstedt is unknown but <strong>the</strong> adults inhabit shady woods.<br />
<strong>An</strong>thomyiidae<br />
(Figs: larvae 855- 902, eggs I 097- <strong>10</strong>99, puparia 1263- 1265, leaf-mines 1309- 131 0)<br />
About I ,200 species in some 55 genera <strong>of</strong> <strong>An</strong>thomyiidae are known <strong>to</strong> science but no<br />
doubt more species await description in this neglected family. The <strong>immature</strong> <strong>stages</strong> are<br />
little known; some are phy<strong>to</strong>phagous and a few species are well known agricultural<br />
pests. O<strong>the</strong>rs live as scavengers in decaying organic matter. Some 220 species in 34<br />
genera occur in Britain. Papers by Dusek (1969, 1970) are useful for <strong>the</strong> identification <strong>of</strong><br />
<strong>the</strong> <strong>immature</strong> <strong>stages</strong>, especially <strong>of</strong> <strong>the</strong> economically important species. O<strong>the</strong>r references<br />
are cited under each genus. The arrangement <strong>of</strong> <strong>the</strong> posterior fleshy processes is a<br />
valuable help for spot identifications among <strong>the</strong> economic species. Genera not mentioned<br />
may be assumed <strong>to</strong> be unknown in <strong>the</strong> <strong>immature</strong> <strong>stages</strong>. Reared adults should be<br />
identified using Hennig ( 1966-1976) who also gives a full list <strong>of</strong> rearing records and<br />
parasites. Much biological information is also included in Stubbs & Chandler ( 1978).<br />
The larvae <strong>of</strong> Chirosia species (figs 855- 857) mine <strong>the</strong> leaves <strong>of</strong> ferns (Pteridium,<br />
Athyriumfilix-femina, Thelypteris palustris), some causing <strong>the</strong> tips <strong>of</strong> <strong>the</strong> frond <strong>to</strong> roll<br />
over (fig. 131 0) (see also Acrostilpna below).<br />
Fucellia species frequent seaweed on seashores. Egglishaw ( 1960c) describes <strong>the</strong> lifehis<strong>to</strong>ry<br />
<strong>of</strong> Fucellia maritima Haliday (figs 858- 862), which he found mostly in <strong>the</strong><br />
smaller wrack banks.<br />
Chias<strong>to</strong>cheta species (figs 863-865) lay <strong>the</strong>ir eggs in <strong>the</strong> unripe seed-heads <strong>of</strong> Trollius<br />
and <strong>the</strong> larvae bore in<strong>to</strong> and feed upon <strong>the</strong> seeds, eventually leaving <strong>the</strong> seed-head <strong>to</strong><br />
pupate in <strong>the</strong> earth.<br />
Pegohylemyia fugax (Meigen) (figs 866-868) is associated mostly with decaying<br />
vegetation. From May <strong>to</strong> Oc<strong>to</strong>ber its eggs are numerous on cruciferous crops, usually<br />
on dead leaves, in cracked and broken stems, etc. Larvae have been recovered from<br />
cauliflower heads, Brussels sprouts, swedes, spring cabbages, cabbage seedlings,<br />
turnips, lettuce, beet and oat seedlings. The larvae <strong>of</strong> P. gnava (Meigen) (figs 869-871)<br />
130
(<strong>the</strong> lettuce seed fly) develop in Lactuca species. P. silvatica (Robineau-Desvoidy) has<br />
been reared from fungi (agarics, boleti, Phallus). P. phrenione (Seguy), P. dissecta<br />
(Meigen) and P. laterella Collin are all associated with <strong>the</strong> grass-fungus Epichloe<br />
typhina Tulasne (though <strong>the</strong>y have been misidentified in <strong>the</strong> literature as <strong>An</strong>thomyia<br />
spreta, e.g. Lucas, 1909; Chor<strong>to</strong>phila humerella, etc.). The larvae <strong>of</strong> P. seneciella<br />
(Meade) and P. jacobaeae (Hardy) develop in <strong>the</strong> flower-heads <strong>of</strong> Senecio and have<br />
been used in Australia and New Zealand in <strong>the</strong> biological control <strong>of</strong> introduced weeds;<br />
P. pseudomaculipes Strobl develops in <strong>the</strong> flower-heads <strong>of</strong> Solidago virgaurea L.; P.<br />
brunneilinea (Zetterstedt) in Centaurea ( base <strong>of</strong> stem); P. sonchi (Hardy) in Sonchus;<br />
P. spinosa (Rondani) in Achillea millefolium (stem); P. sanctimarci (Czerny) in Allium<br />
ursinum.<br />
Lasiomma meadei (Kowarz) has been reared from cow and human dung and birds'<br />
nests. L. anthomyinum (Rondani) and L. oc<strong>to</strong>guttatum (Zetterstedt) have also been<br />
reared from birds' nests (Collin, 1939; Hicks, 1959). Beaver ( 1969) has reared <strong>the</strong> latter<br />
from dead snails and a puparium has been described from an 'alluvial deposit <strong>of</strong> River<br />
Rhine near Herwen' by S<strong>to</strong>rk (1936). L. melania and L. infrequens (both described by<br />
Ackland ( 1965)) may feed on <strong>the</strong> seeds <strong>of</strong> larch as do related species abroad.<br />
Hydrophoria lancifer (Harris) ( = conica (Wiedemann)) breeds in human dung. There<br />
is a specimen <strong>of</strong> H. linogrisea Meigen in <strong>the</strong> <strong>British</strong> Museum (Nat. Hist.) labelled<br />
' round <strong>An</strong>drena burrows', but no association between Hydrophoria and bees is known.<br />
Larvae <strong>of</strong> Acrostilpna latipennis (Zetterstedt) occur in <strong>the</strong> leaf-stem <strong>of</strong> <strong>the</strong> fern,<br />
Athyriumfilix-femina.<br />
According <strong>to</strong> S<strong>to</strong>rk ( 1936) larvae <strong>of</strong> Craspedochoeta pullula (Zetterstedt) have been<br />
recorded as injurious <strong>to</strong> Iris and Gladiolus by eating <strong>the</strong> flowers, leaves and stems, but<br />
I know <strong>of</strong> no recent (or <strong>British</strong>) records <strong>of</strong> this. Hammond & Smith ( 1953) record a<br />
specimen <strong>of</strong> C. pullula which had emerged in a tin containing seedling lettuce leaves<br />
and sterilized peat and leaf-mould provided as pabulum for a caterpillar <strong>of</strong> Agrotis<br />
segetum (Lep., Noctuidae). The caterpillar died and parasitism was thus suspected<br />
but seemed hardly likely. However <strong>the</strong>re are two specimens <strong>of</strong> C. cannabina Stein in<br />
<strong>the</strong> <strong>British</strong> Museum (Nat. Hist.) ex coli. W. Rait-Smith (a Lepidopterist but also a<br />
careful recorder <strong>of</strong> <strong>the</strong>ir parasites, e.g. see Morley & Rait-Smith, 1933), labelled 'Dip.<br />
parasite <strong>of</strong> Tinea lapella Huhner'. At least we were not alone in our mistake if such<br />
it proves <strong>to</strong> be (see also comments under rearing Tachinidae; and Smith, 1974b).<br />
These records seem doubtful or possibly chance occurrences. Both C. cannabina and<br />
T. lapella occur in birds' nests! Beaver ( 1969) has reared C. pull uta from dead<br />
snails (pupating in <strong>the</strong> soil) and suggests that it probably breeds in decaying animal<br />
matter.<br />
<strong>An</strong>thomyia species have been reared from fungi: A. pluvialis (L.) from several agarics,<br />
boleti and Phallus; A. procellaris Rondani (figs 872- 874) from Pleurotus ostreatus<br />
Quelet. Both species have also been reared from birds' nests (Hicks, 1959- 1971), where<br />
Keilin ( 1924b) claims <strong>the</strong>y feed on 'various decomposed organic substances including<br />
greasy scales and down left by <strong>the</strong> birds'.<br />
The larvae <strong>of</strong> Phorbia securis (Tiensuu) ( =genitalis auctt.) (figs 875- 877) and<br />
P. sepia (Meigen) (late-wheat shoot-flies) attach <strong>the</strong> shoots <strong>of</strong> wheat and occasionally<br />
o<strong>the</strong>r cereals.<br />
The larvae <strong>of</strong> some Leucophora species live upon <strong>the</strong> pollen masses s<strong>to</strong>red by bees or<br />
have o<strong>the</strong>rwise been associated with Aculeate Hymenoptera as follows: L. grisella<br />
Hennig (Panurgus calcaratus (Scopoli), Cerceris arenaria (L.)); L. cinerea Robineau<br />
Desvoidy (Ha/ictus nitidiusculus (Kirby); L. obtusa (Zetterstedt) (<strong>An</strong>drenafulva (Muller<br />
in Allioni)); L. personata (Collin) (<strong>An</strong>drena labia/is (Kirby), A. nigroanenea (Kirby)<br />
and A. trimmerana (Kirby)); L. sericea Robineau-Desvoidy (<strong>An</strong>drena haemorrhoa (F.)<br />
(=albicans misident), A.fulva) (Collin, 1921 ).<br />
Eustalomyia species (figs 878) are similarly associated with Aculeate Hymenoptera<br />
(Smith, 1971), especially <strong>the</strong> burrows <strong>of</strong>Sphecidae.<br />
131
The genus Delia ( = Erioischia, Lep<strong>to</strong>hylemyia) contains several species which are<br />
well known agricultural and horticultural pests. D. antiqua (Meigen) (onion fly) (figs<br />
879- 881) attacks young onion plants, leeks and shallots in gardens and market gardens<br />
especially in <strong>the</strong> Midlands and East <strong>An</strong>glia; it kills young plants outright and also<br />
tunnels in onion and tulip bulbs. D. radicum (L.) ( = brassicae (H<strong>of</strong>fmannsegg in<br />
Wiedemann)) (cabbage root fly) (figs 882-886) inflicts most damage on early cauliflowers<br />
and summer cabbages but radishes, turnips and swedes are also attacked. The<br />
larvae are mostly root feeders but also tunnel <strong>the</strong> stems. The larvae <strong>of</strong> D. cardui<br />
(Meigen) (carnation fly) attack carnations and sweet williams, first mining <strong>the</strong> leaves<br />
<strong>the</strong>n later burrowing down in<strong>to</strong> <strong>the</strong> stem and devouring <strong>the</strong> pith. D. coarctata (Fallen)<br />
(wheat bulb fly) (figs 887- 890) attacks winter wheat, barley and rye; its known wild host<br />
is couch grass (Agropyron repens (L.)). The larvae in <strong>the</strong> central shoot kill <strong>the</strong> growing<br />
point, <strong>the</strong>n move <strong>to</strong> ano<strong>the</strong>r plant. D. echinata (Seguy) (spinach stem fly) (figs 891- 893)<br />
attacks seedlings <strong>of</strong> spinach forming small blotch mines in <strong>the</strong> leaves, <strong>the</strong>n tunnelling<br />
via <strong>the</strong> veins or midribs in<strong>to</strong> <strong>the</strong> stem (Miles, 1953). D.floralis (Fallen) (turnip root fly)<br />
attacks turnips, especially in Scotland, making wounds which admit pathogens causing<br />
<strong>the</strong> swollen roots <strong>to</strong> rot; it may also cause severe damage <strong>to</strong> seedlings and newly<br />
transplanted crops <strong>of</strong> onions and leeks (Miles, 1953). D. platura (Meigen) ( = cilicrura<br />
Rondani) (bean seed fly) (figs 894-897) is widely polyphagous and attacks French and<br />
runner beans, broad beans, onions, Brassica, lettuce, and occasionally winter cereals.<br />
O<strong>the</strong>r known associations <strong>of</strong> Delia species with higher plants (not all are <strong>British</strong> rearing<br />
records) are as follows: D. coronariae (Hendel), Lychnis flos-cuculi L. (stem);<br />
D. flavifrons (Zetterstedt), Silene cucubalus Wibel (seed); D. florilega (Zetterstedt)<br />
( = trichodacty la Rondani), Raphanus sativus L., Armoracia rusticana Gaertner,<br />
Brassica spp., Cichorium intybus L. (root); D. planipalpis (Stein), Brassica napus L., B.<br />
rapa L. (in turnip roots); D. quadripila (Stein), Honckenya pep/aides (L.) (leaf-miner).<br />
Fur<strong>the</strong>r information on this important genus is given in Hennig (1966-1976) and<br />
Stubbs & Chandler (1978). Some species <strong>of</strong> Delia have been reared from fungi as<br />
follows: D. a/hula (Fallen), from Psathyrella; D. antiqua, several agarics (Russula ,<br />
Amanita, Tricholoma , Armillaria) but <strong>the</strong> precise association with fungi requires investigation,<br />
see above); D. frontella (Zetterstedt) from Suillus bovinus (L.); <strong>the</strong> widely<br />
polyphagous D. platura (see also above) from young plasmodium <strong>of</strong> Fuligo.<br />
Subhylemyia longula (Fallen) has been reared from dead snails (Cepaea nemoralis<br />
(L.)); it pupates in <strong>the</strong> soil (Beaver, 1969).<br />
The larvae <strong>of</strong> some species <strong>of</strong> Hylemya occur in cow dung as follows: H. nigrimana<br />
(Meigen), H. vagans (Panzer) ( =strigosa (F.)) and H. variata (Fallen). Adults <strong>of</strong><br />
<strong>the</strong>se and o<strong>the</strong>r species have been seen on dung (including human) but not reared<br />
from it. H. latifrons Schnabl & Dziedzicki has been reared from fungi (Boletus<br />
sub<strong>to</strong>men<strong>to</strong>sus L.).<br />
Heterostylodes species have been reared from flower-heads <strong>of</strong> Compositae<br />
(Hieracium, Pier is, Leon<strong>to</strong>don and Hypochoeris). Paregle audacula (Harris) ( = radicum<br />
<strong>of</strong> authors) breeds in cow, human and dog dung; P. cinerella (Fallen) is recorded from<br />
<strong>the</strong> dung <strong>of</strong> hedgehog, dog and cow/horse manure. The larvae <strong>of</strong> all Egle species<br />
develop in <strong>the</strong> catkins <strong>of</strong> Salix and Populus tremula L.<br />
Caly<strong>the</strong>a nigricans (Robineau-Desvoidy) occurs in pig dung and Nupedia aestiva<br />
(Meigen) breeds in cow dung but <strong>the</strong> life-his<strong>to</strong>ries <strong>of</strong> <strong>the</strong> o<strong>the</strong>r common species in <strong>the</strong>se<br />
two genera appear <strong>to</strong> be little known. Paradelia palliceps (Zetterstedt) ( = Pseudonupedia<br />
setinerva Ringdahl) has been reared from fungi (Suillus granulatus (L.)), and<br />
Emmesomyia species probably breed in excrement; adults have been seen on human<br />
and horse dung.<br />
The habits <strong>of</strong> <strong>the</strong> large genus Pegomya (42 <strong>British</strong> species) are about evenly divided<br />
between feeding on fungi and <strong>the</strong> higher plants. Full host lists are provided by Hennig<br />
(1966-1976) and Stubbs & Chandler (1978), but here <strong>the</strong> fungus feeders are listed first<br />
and selected plant feeders are treated below.<br />
132
Fungus feeding Pegomya: calyptrata (Zetterstedt), deprimata (Zetterstedt),fulgens<br />
(Meigen), furva Ringdahl, geniculata Bouche, incisiva Stein, macula/a Stein,<br />
pal/idoscutel/ata (Zetterstedt), pulchripes (Loew), rufina (Fallen), tabida (Meigen),<br />
tenera (Zetterstedt), transversa (Fallen), ulmaria Rondani, vittigera (Zetterstedt) and<br />
:::onata (Zetterstedt).<br />
The larvae <strong>of</strong> Pegomya hyoscami (Panzer) (beet or mangold fly or beet leaf miner)<br />
(figs 898- 902) form blister mines on beet and mangolds. Most damage occurs on late<br />
sown plants which have little leaf area developed when <strong>the</strong> first brood <strong>of</strong> flies appears.<br />
P. rubivora (Coquillett) (loganberry cane fly) attacks stems <strong>of</strong> Rubus idaeus L. and<br />
Filipendula.<br />
Mycophaga testacea Gimmerthal has been reported as having carnivorous larvae in<br />
many fungi but this seems very doubtful. Adults occur near fungi and may have been<br />
confused with similar adult Muscidae which do have carnivorous larvae (see also<br />
Muscidae).<br />
Fanniidae<br />
(Figs: larvae 903- 908, egg 1<strong>10</strong>0, puparium 1266)<br />
There are some 265 species <strong>of</strong> Fanniidae contained in 4 genera. The family is best<br />
represented in <strong>the</strong> Palaearctic (<strong>10</strong>7 species) and Nearctic (<strong>10</strong>5, <strong>of</strong> which 29 are<br />
Holarctic) regions. There are 59 <strong>British</strong> species representing two genera. Larval<br />
Fanniidae are very characteristic with a thickened coarse cuticle and <strong>the</strong> body is dorsoventrally<br />
flattened with a number <strong>of</strong> fleshy dorsal and lateral processes, which may be<br />
plumed, pubescent or simple. The eggs have flange-like processes (fig. 1<strong>10</strong>0). The<br />
<strong>immature</strong> <strong>stages</strong> develop in a wide range <strong>of</strong> decaying organic matter usually <strong>of</strong><br />
vegetable origin but also in excrement and in carrion. Some occur in fungi or <strong>the</strong><br />
burrows and nests <strong>of</strong> mammals, birds and insects (social Hymenoptera) and a few are<br />
<strong>of</strong> medical or veterinary importance in cases <strong>of</strong> myiasis. The larvae <strong>of</strong> only 16 <strong>of</strong> <strong>the</strong><br />
<strong>British</strong> species are known (Lyneborg, 1970) but rearing data for a fur<strong>the</strong>r 12 species are<br />
available.<br />
Piezura species have been reared from fungi; P. bole<strong>to</strong>rum (Rondani) from<br />
an unidentified fungus and P. graminicola (Zetterstedt) from Coprinus micaceus<br />
Fries.<br />
Fannia aequi/ineata Ringdahl has been reared from a blackbird's nest, wood detritus<br />
and <strong>the</strong> fungi lnonotus dryadeus and Daldinia concentrica; <strong>the</strong> adults are associated with<br />
sap runs. O<strong>the</strong>r fungus-feeding Fannia include F. canicularis (L.) (many decaying fungi<br />
and o<strong>the</strong>r media, see below), F. diffici/is (Stein) (Lactarius. Boletus), F. immutica Collin<br />
(indet. fungus), F. incisurata (Zetterstedt) (indet. fungus), F. manicata (Meigen)<br />
(Amanita, Laetiporus, boleti), F. melania (Dufour) (several boleti, Armi//aria,<br />
Lactarius, Tricholoma, Phallus), F. moni/is (Haliday) (Pieurotus, Polyporus), F. lepida<br />
(Wiedemann) ( =mutica (Zetterstedt)) (indet. fungus), F. scalaris (F.) (Boletus,<br />
Polyporus, Laetiporus, o<strong>the</strong>r media, see below).<br />
Species reared from birds' nests (hosts <strong>of</strong> <strong>the</strong> apprarently more specific species are<br />
cited) include: F. canicularis, F. c/ara Collin (little owl, heron), F. coracina (Loew), F.<br />
difficilis, F. hirundinis Ringdahl (sand martins), F. incisurata, F. manicata, F. monilis,<br />
F. nidica Collin, F. parva (Stein) (house martin), F. rondanii Strobl (house martin),<br />
F. scalaris, F. vesperti/ionis Ringdahl (starling), F. /ineata Stein (heron).<br />
Fannia canicularis, F. fuscula (Fallen) (fig. 908), F. coracina, F. scalaris and<br />
F. vesparia (Meade) have all been reared from <strong>the</strong> nests <strong>of</strong> wasps or bees.<br />
F. vesperti/ionis has been reared from droppings <strong>of</strong> <strong>the</strong> noctule bat and F. canicularis<br />
has also been associated with bats.<br />
Some o<strong>the</strong>r species have been reared from wood detritus, litter or woodland soil, e.g.<br />
F. gotlandica Ringdahl, F. umbrosa (Stein), F. minutipalpis Stein, F. polychaeta (Stein),<br />
133
F. postica (Stein), and F. speciosa (Villeneuve). F. genua/is (Stein) has been found in leaf<br />
litter.<br />
Clearly <strong>the</strong> examination <strong>of</strong> fungi, old nests and <strong>the</strong> droppings <strong>of</strong> smaller mammals<br />
(especially bats) yields <strong>the</strong> rarer species <strong>of</strong> Fannia and should help elucidate our<br />
knowledge <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ries and <strong>immature</strong> <strong>stages</strong>.<br />
Some <strong>of</strong> <strong>the</strong> commoner species <strong>of</strong> Fannia mentioned above have a wide range <strong>of</strong><br />
habits which may bring <strong>the</strong>m in<strong>to</strong> a closer involvement with man and give <strong>the</strong>m a<br />
considerable medical importance. Fannia canicularis (figs 903- 905) is <strong>the</strong> lesser housefly,<br />
a common domestic species with a cosmopolitan distribution. The larvae develop<br />
in a variety <strong>of</strong> media including rotting vegetables and apples, large accumulations <strong>of</strong><br />
faeces (e.g. on poultry farms) and in carrion. F. scalaris (fig. 907), <strong>the</strong> latrine fly,<br />
is essentially an outdoor species associated with primitive lava<strong>to</strong>ries and cesspits.<br />
Optimum conditions occur in semi-liquid masses <strong>of</strong> faeces, especially pigs', but also <strong>of</strong><br />
o<strong>the</strong>r animals, including man. Both species have been involved in cases <strong>of</strong> urogenital<br />
and intestinal myiasis and may occur in urine soaked babies' napkins and cot blankets<br />
(Zumpt, 1965). These species may also have an importance in forensic investigations<br />
(Smith, 1986b ). F. manicata (fig. 906) has also been implicated in myiasis and forensic<br />
cases but is normally found in decaying vegetable matter such as garden refuse, putrid<br />
cabbage leaves, etc. and has also been reared from rabbit excrement and a dead gull.<br />
Muscidae<br />
(Figs: larvae 909-966, eggs 1<strong>10</strong>1- 1<strong>10</strong>8, cover, puparia 1267-1281, cocoon 1272)<br />
There are some 3,900 species <strong>of</strong> Muscidae in some 170 genera and <strong>the</strong> family occurs<br />
abundantly in all <strong>the</strong> major zoogeographical regions. Six subfamilies are usually<br />
recognised and 5 <strong>of</strong> <strong>the</strong>se, including 280 species in 44 genera, occur in Britain.<br />
Larval Muscidae develop in a wide range <strong>of</strong> substances from decaying vegetable<br />
matter, wood, fungi and living plants, <strong>the</strong> nests and burrows <strong>of</strong> birds, mammals and<br />
insects, <strong>to</strong> rot-holes, water margins and running water. A few species cause myiasis in<br />
man and animals. Many larvae are carnivorous at least in <strong>the</strong> later instars and associated<br />
with this habit is a longer incubation period in <strong>the</strong> egg which may hatch as second<br />
or even third ins tar larvae. The females <strong>of</strong> some species deposit third stage larvae. Some<br />
species form cocoons (fig. 1272) prior <strong>to</strong> pupariation (Ferrar, 1980).<br />
Skidmore (1985) describes, keys and illustrates <strong>the</strong> known larvae <strong>of</strong> Muscidae,<br />
amounting <strong>to</strong> 440 <strong>of</strong> <strong>the</strong> world species. He defines <strong>the</strong>m according <strong>to</strong> <strong>the</strong>ir habits as<br />
trimorphic (3 instars) saprophages, facultative carnivores, <strong>to</strong> monomorphic (one<br />
instar, <strong>the</strong> last) carnivores.<br />
The relationship between <strong>the</strong> morphology and biology <strong>of</strong> <strong>the</strong> larvae and puparia <strong>of</strong><br />
Muscidae may be summarised (after Skidmore, 1985) as follows (figure numbers refer<br />
<strong>to</strong> <strong>the</strong> present Handbook):<br />
I. A sieving mechanism (fig. 911, sm) occurs in all known trimorphic larvae and in<br />
some dimorphic ones. Micro-sectioning <strong>of</strong> <strong>the</strong> pharyngeal floor <strong>of</strong> a wide range <strong>of</strong><br />
monomorphic and dimorphic forms is desirable <strong>to</strong> ascertain if <strong>the</strong>se structures occur<br />
widely. The structure is described in detail by Keilin ( 1917).<br />
2. All known carnivorous species have well-developed oral bars (fig. 911 , ob) and<br />
anterior ribbons (axillary rods) (fig. 911, ar), but in some species <strong>the</strong>se are apparently<br />
being lost due <strong>to</strong> reversal <strong>to</strong> non-carnvivorous diet.<br />
3. Massive posterior spiracles indicate trimorphism.<br />
4. Minute posterior spiracles indicate reduced number <strong>of</strong> ins tars, <strong>the</strong> smallest being<br />
probably monomorphic.<br />
5. Obligative carnivores found in dung do not have gut discoloured by <strong>the</strong> dung.<br />
6. Dung-frequenting larvae with dung in gut are coprophagous or facultative<br />
carnivores.<br />
134
7. Most wholly aquatic larvae have false legs and <strong>of</strong>ten long anal spiracular<br />
processes.<br />
8. Obligate phy<strong>to</strong>phages have oral bars (fig. 911 , ob), <strong>the</strong> hypopharyngeal sclerite<br />
(fig. 911, hs) massively enlarged and <strong>the</strong> pharyngeal sclerite (fig. 911, ps) <strong>of</strong>ten fused <strong>to</strong><br />
it.<br />
O<strong>the</strong>r useful works are Keilin ( 1917), Keilin & Tate ( 1930) and Skidmore ( 1973).<br />
Larvae <strong>of</strong> <strong>the</strong> subfamilies <strong>of</strong> Muscidae occurring in Britain may be separated in <strong>the</strong><br />
third instar as follows (simplified after Skidmore ( 1985) who recognises I 0 subfamilies<br />
(8 <strong>British</strong>)).<br />
Pharyngeal sclerite (ps) <strong>of</strong>mouthparts with a distinct atrial angle (at) (fig. 919) 2<br />
Atrial angle absent or if present (some Spilogona, Phaonia) <strong>the</strong>n weak and obtuse 5<br />
2 Posterior spiracles with slits straight, sinuate or curved; parallel convergent or radiate (figs<br />
925, 930) (never serpentine or <strong>to</strong>rtuous) . . . . . . . . . . . . . . 3<br />
Posterior spiracular slits serpentine <strong>to</strong> <strong>to</strong>rtuous, peripheral or encircling (figs 9<strong>10</strong>, 913, 916) 4<br />
3 Posterior spiracles on dorsal surface <strong>of</strong> last segment; in wasps' nests . Achanthipterinae<br />
Posterior spiracles o<strong>the</strong>rwise; usually in dung, carrion or soil . . . . . . Azeliinae<br />
4 Lower half (ventral cornu, vc) <strong>of</strong> pharyngeal sclerite (ps) with a strong '<strong>to</strong>oth' (posterior<br />
projection, pp)(figs 788,919, pp) . . . . . . . . . . . . . Muscinae<br />
At most only a small weak pharyngeal <strong>to</strong>oth present (figs 958, 962, 966) . S<strong>to</strong>moxyinae<br />
5 Posterior spiracles with slits convergent, straight <strong>to</strong> <strong>to</strong>rtuous (fig. 934). Mouthparts with<br />
sieving mechanisms (fig. 911, sm). Body never with false legs or caudal processes . . .<br />
. . . . . . . . . . . . . . . . . . . . . . . Reinwardtiinae<br />
Slits <strong>of</strong> posterior spiracles parallel <strong>to</strong> radiate, straight <strong>to</strong> curved or angular, never <strong>to</strong>rtuous<br />
(figs 936, 940). Body sometimes with long false legs or caudal processes (e.g. Limnophora)<br />
(figs 938, 949) . . . . . Mydaeinae, Coenosiinae, Phaoniinae<br />
The classification used mainly follows Skidmore ( 1985) but as far as possible follows<br />
<strong>the</strong> sequence <strong>of</strong> genera in <strong>the</strong> Check List (Smith et al. , 1976). For nomenclature see Pont<br />
(in So6s & Papp, 1986).<br />
Achanthipterinae. The larvae <strong>of</strong> Achanthiptera rohre/liformis (Robineau-Desvoidy)<br />
(fig. 909) feed on decaying matter in wasps' nests ( Vespula).<br />
Muscinae. The larvae <strong>of</strong> this subfamily are trimorphic and saprophagous or<br />
(Mesembrina) faculatative carnivores.<br />
Polietes species (figs 9<strong>10</strong>-912) are found in cow dung or (P. domi<strong>to</strong>r (Harris)<br />
( = albolineata Fallen) and P. steinii Ringdahl) in horse dung. They become carnivorous<br />
in <strong>the</strong> third instar, though <strong>the</strong>re is no evidence <strong>of</strong> this yet for P. lardaria (F.).<br />
Mesembrina meridiana (L.) (figs 913-915) breeds in cow dung and less frequently<br />
in horse dung. Occasionally <strong>the</strong> first stage is spent entirely within <strong>the</strong> egg and a<br />
second instar larva hatches. Even normally development is very rapid and within 24<br />
hours <strong>of</strong> hatching from <strong>the</strong> egg <strong>the</strong> larva will reach <strong>the</strong> third instar and <strong>the</strong> whole<br />
life-his<strong>to</strong>ry takes only a week in summer. Two adults in <strong>the</strong> B.M. (N.H.) are labelled<br />
'bred from half-grown larvae which destroyed 40 larvae, large and small, <strong>of</strong> Musca<br />
corvina'.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> <strong>the</strong> <strong>British</strong> Pyrellia are unknown, but <strong>the</strong> only known larval<br />
pabulum is <strong>the</strong> dung <strong>of</strong> cattle and horses.<br />
Eudasyphora cyane/la (Meigen) and E. cyanicolor (Zetterstedt) have been reared<br />
from sheep dung. Neomyia ( = Or<strong>the</strong>/lia) and More/lia species breed mostly in cow<br />
dung.<br />
Musca domestica L. (figs 916--919) is <strong>the</strong> ubiqui<strong>to</strong>us house fly, a truly synanthropic<br />
species that has followed man around <strong>the</strong> world, breeds in his refuse and is a major<br />
mechanical transmitter <strong>of</strong> disease organisms in <strong>the</strong> adult stage. Up <strong>to</strong> 2,500 eggs may be<br />
laid and as many as 18 broods a year may be reared. It is perhaps fortunate that many<br />
135
predaceous muscid larvae (Muscina, Myospila, Hydrotaea) attack <strong>the</strong> larvae <strong>of</strong> M.<br />
domestica (and o<strong>the</strong>r Musca species) and that <strong>the</strong> fungus En<strong>to</strong>mophthora muscae Cohn<br />
also plays a part in controlling its numbers. In natural conditions in rural areas eggs are<br />
laid on horse dung but human, cow and poultry dung or material contaminated with<br />
excrement, decaying vegetable matter, garbage, decomposing foodstuffs, meat and<br />
carcases may all provide suitable food for <strong>the</strong> larvae, which can also complete <strong>the</strong>ir<br />
development on urine alone. M. domestica is one <strong>of</strong> <strong>the</strong> few Diptera <strong>to</strong> have an individual<br />
monograph (West, 1951) and a bibliography (West & Peters, 1973) devoted <strong>to</strong> it;<br />
Bollwig (1946) gives a detailed study <strong>of</strong> <strong>the</strong> larval sense organs. Musca autumnalis<br />
De Geer is known as <strong>the</strong> face fly because adults cause irritation <strong>to</strong> cattle by feeding<br />
on <strong>the</strong>ir body secretions, especially around <strong>the</strong> eyes and muzzle. They are also vec<strong>to</strong>rs<br />
<strong>of</strong> Thelazia (parasitic eye worm). The larvae develop in cow and horse dung and<br />
occasionally in carrion.<br />
Azeliinae. The larvae <strong>of</strong> this subfamily are trimorphic facultative <strong>to</strong> dimorphic (or<br />
monomorphic) obligate carnivores.<br />
The larvae <strong>of</strong> Azelia (figs 920-921) occur in horse and cow dung, where, in <strong>the</strong>ir final<br />
instar, <strong>the</strong>y prey upon small nema<strong>to</strong>cerous and acalyptrate larvae.<br />
Thricops larvae have been found in <strong>the</strong> soil under logs, leaf litter and wet moss where<br />
<strong>the</strong>y are presumed <strong>to</strong> be carnivorous.<br />
Alloeostylus larvae have been found in decayed logs and have minute posterior<br />
spiracles which suggests that <strong>the</strong>y hatch in <strong>the</strong> final instar and are probably obligate<br />
carnivores (Skidmore, 1985).<br />
Larvae <strong>of</strong> Drymeia ( = Trichopticoides, Pogonomyia) species are probably soil<br />
dwellers in deep leaf litter in woodland but D. vicana (Harris) ( = decolor Fallen) (fig.<br />
922) occurs in old cow pats and is probably an obligate carnivore.<br />
The larva <strong>of</strong> Potamia ( = Dendrophaonia) lit<strong>to</strong>ralis Robineau-Desvoidy ( = querceti<br />
Bouche) (figs 923-924) is a facultative carnivore and has been reared from bat droppings,<br />
rabbit dung, human faeces, rotten wood, nests <strong>of</strong> birds and social insects, liquid<br />
substrates in rubbish dumps, cess pits, privies and fungi.<br />
Ophyra capensis (Wiedemann) (fig. 1270) has been reared from human faeces,<br />
carrion, dead locusts, and <strong>the</strong> nests <strong>of</strong> birds and mammals. This species was rare until<br />
recently, but changed methods <strong>of</strong> poultry farming favour its development and it is now<br />
common in poultry battery farms where it breeds with 0. ignava, Musca domestica,<br />
Muscina stabulans and Fannia canicularis (Conway, 1970). 0. ignava (Harris)<br />
( = leucos<strong>to</strong>ma Wiedemann) (figs 925-928) breeds in dung, carrion, birds' nests and<br />
rotting foodstuffs. It appears <strong>to</strong> be strongly attracted <strong>to</strong> ammonia resulting from<br />
decaying matter. Ophyra is <strong>of</strong>ten <strong>of</strong> value in forensic investigations, especially when<br />
corpses have been concealed and not <strong>the</strong>refore exposed <strong>to</strong> <strong>the</strong> normal blowfly fauna<br />
(Smith, 1986b ).<br />
Hydrotaea species (adults <strong>of</strong> some species are sweat flies) lay relatively few eggs and<br />
<strong>the</strong> larvae usually hatch in <strong>the</strong> second instar which lasts only a few hours. They are<br />
carnivorous in <strong>the</strong> third instar. The majority <strong>of</strong> species breed only in <strong>the</strong> dung <strong>of</strong><br />
herbivores or in humus soil. H. basdeni Collin and H. nidicola Malloch are known only<br />
from birds' nests and H. floccosa Macquart ( = armipes <strong>of</strong> authors) has been reared<br />
from rotting compost, seaweed, sheep carcases, dead snails and fungi. Graham-Smith<br />
(1916) claims that H. dentipes (F.) (figs 929-931) is <strong>the</strong> most important species with<br />
carnivorous larvae and destroys enormous numbers <strong>of</strong>larvae <strong>of</strong> o<strong>the</strong>r flies . It occurs on<br />
carrion and in fungi and so does H. armipes (Fallen) (=occult a (Meigen)); both<br />
species are occasionally <strong>of</strong> forensic importance (Smith, 1986b ). Females <strong>of</strong> H. irritans<br />
(Fallen) (fig. 932) (<strong>the</strong> sheep head fly) are a considerable nuisance <strong>to</strong> man, cattle and<br />
sheep in late autumn. The species was formerly thought <strong>to</strong> belong <strong>to</strong> <strong>the</strong> true cow dung<br />
community but is now known <strong>to</strong> develop in mostly pasture soil under long grass or<br />
along <strong>the</strong> edges <strong>of</strong> woodland where <strong>the</strong> density <strong>of</strong> mature larvae may be c. 15,000 per<br />
136
acre (Robinson, J. & Luff, 1976; Skidmore, 1986). Lobanov (1970a) provides a useful<br />
paper on Hydrotaea.<br />
Reinwardtiinae. Muscina levida (Harris) ( = assimilis Fallen) develops in a wide range<br />
<strong>of</strong> fungi; it also occurs in <strong>the</strong> excrement <strong>of</strong> man and domestic animals. Muscina prolapsa<br />
(Harris) ( = pabulorum Fallen) develops in carrion, especially <strong>of</strong> larger carcases, where<br />
it is saprophagous until <strong>the</strong> third instar when it becomes a facultative carnivore. The<br />
larvae <strong>of</strong> M. stabulans (Fallen) (figs 933- 934) are found in rotting fungi, fruits, broken<br />
eggs, excrement and carrion where <strong>the</strong>y become carnivorous and even cannibalistic in<br />
<strong>the</strong> later instars; in birds' nests <strong>the</strong>y will attack and kill nestlings and have been found in<br />
<strong>the</strong> dead bodies <strong>of</strong> o<strong>the</strong>r insects. All three species have been reared from nests <strong>of</strong> <strong>the</strong><br />
wasp Vespula vulgaris (L.).<br />
Phaoniinae. The larvae <strong>of</strong> this subfamily are dimorphic or monomorphic obligate<br />
carnivores.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Lophosceles are unknown.<br />
Phaonia is <strong>the</strong> largest <strong>British</strong> genus <strong>of</strong> Muscidae with 45 species. Phaonia<br />
( = Dialy tina) atriceps (Loew) occurs under <strong>the</strong> loose outer sheaths and in <strong>the</strong> workings<br />
<strong>of</strong> o<strong>the</strong>r insects in <strong>the</strong> plants Typha latifolia L., Phragmites, etc. Most Phaonia species<br />
appear <strong>to</strong> hatch from <strong>the</strong> egg in <strong>the</strong> third instar as obligate carnivores. Development<br />
occurs in a wide range <strong>of</strong> habitats but groups <strong>of</strong> species appear <strong>to</strong> favour particular<br />
media (Skidmore, 1985). P. exoleta (Meigen) (figs 935- 938) develops in water holes in<br />
old foliferous trees ( Ulmus, Fraxinus, Aesculus, Acer); its larva swims actively and feeds<br />
voraciously on culicid larvae. A single P. exoleta larva may consume <strong>10</strong>0 mosqui<strong>to</strong><br />
larvae. Keilin ( 1917) and Tate (1935) describe three groups <strong>of</strong> long mobile hairs on <strong>the</strong><br />
ventral surface <strong>of</strong> <strong>the</strong> thoracic segments but <strong>the</strong>se appear <strong>to</strong> be absent in <strong>the</strong> specimen <strong>of</strong><br />
P. exoleta (in <strong>the</strong> B.M. (N.H.)) illustrated by Oldroyd (1964). I have combined <strong>the</strong><br />
features <strong>of</strong>Oidroyd's and Tate's illustrations <strong>to</strong> include <strong>the</strong>se hairs (fig. 938). The larva<br />
<strong>of</strong> P. gobertii (Mik) is found under sodden bark (especially on Ulmus, Populus), where it<br />
preys on lonchaeid and clusiid larvae; it also develops in woodland leaf-litter and fungi.<br />
P.laeta (Fallen) (=trigona/is (Meigen), laetabilis Collin) has been reared from tree rotholes.<br />
P. palpata (Stein) has been reared from dead trees. The following Phaonia species<br />
have been reared from fungi: P. canescens Stein (also under bark and in rotten wood<br />
etc.), P. gobertii, P. pallida (F.), P. rufiventris (Scopoli) (=populi Meigen) (and dead or<br />
sickly trees), P. subventa (Harris) ( = variegata Meigen) (also rotten trees, under bark,<br />
dung, carrion and compost).<br />
Species reared from humus soil, under moss, etc include P. valida (Harris) ( = viarum<br />
Robineau-Desvoidy), P. errans (Meigen) (also dung and compost), P. tuguriorum<br />
(Scopoli) (=signata Meigen), P. angelicae (Scopoli) (=basalis Zetterstedt), P. incana<br />
(Wiedemann). P. villana Robineau-Desvoidy ( =mystica <strong>of</strong> authors) has been reared<br />
from dense cushions <strong>of</strong> moss on soil, rocks or tree trunks in woods on calcareous basic<br />
igneous rocks. P. cincta (Zetterstedt) develops in sap runs <strong>of</strong> broad-leaved trees<br />
(especially Ulmus) where larvae prey upon those <strong>of</strong>lonchaeids, clusiids and Myce<strong>to</strong>bia<br />
pallipes Meigen (<strong>An</strong>isopodidae). P. trimaculata (Bouche) (fig. 939) is primarily associated<br />
with herbaceous plants (especially larger Cruciferae) where its larvae prey upon<br />
those <strong>of</strong> various anthomyiids (including Delia brassicae) and Coleoptera.<br />
The genus Helina has 38 <strong>British</strong> species which mostly develop in moss <strong>of</strong> humus soil,<br />
though some may invade adjacent materials (e.g. cow dung). Variations from this habit<br />
are mentioned below. They are obligate monomorphic carnivores. H. quadrum (F.)<br />
appears <strong>to</strong> be predaceous on larvae <strong>of</strong> <strong>the</strong> beetle Phyllopertha horticola (L.) (<strong>the</strong> garden<br />
chafer) in grassland (Smith, 1983). The larvae <strong>of</strong> H. pertusa (Meigen) (figs 940-941) are<br />
found behind loose bark on dead or dying trees where <strong>the</strong>y prey on <strong>the</strong> larvae <strong>of</strong><br />
Lonchaeidae, Clusiidae, Myce<strong>to</strong>philidae, Sciaridae and Cera<strong>to</strong>pogonidae; it is also<br />
found in birds' nests. The larva spins a frail silken cocoon which incorporates particles<br />
137
<strong>of</strong> debris. Larvae <strong>of</strong> H. vicina (Czerny) have been found under moss on tree trunks and<br />
rocks in hilly areas and I found larvae <strong>of</strong> H. reversio (Harris) ( = duplicata Meigen)<br />
under moss on <strong>the</strong> Wrekin in Shropshire (Smith, 1954). The latter species has also been<br />
recorded from a hornets' nest and a tree stump in Denmark (Michelsen, 1977). The<br />
female <strong>of</strong> H. protuberans (Zetterstedt) oviposits in sand dunes and has an oviposi<strong>to</strong>r<br />
reminiscent <strong>of</strong> Asilidae but <strong>the</strong> larval habits and prey are unknown. H . sexmaculata<br />
(Preyssler) ( = punctata Robineau-Desvoidy) has been reared from a swallow's nest and<br />
H. pulchella Ringdahl from <strong>the</strong> nest <strong>of</strong> a tawny owl (Smith, 1960).<br />
Brontaea humilis Zetterstedt has been reared from horse dung.<br />
Mydaeinae. Larvae <strong>of</strong> this subfamily are dimorphic or monomorphic obligate<br />
carnivores. The eggs (fig. 1<strong>10</strong>5) have respira<strong>to</strong>ry horns <strong>of</strong> varying length.<br />
Hebecnema species develop in humus soil, and cow and horse dung; H. umbratica<br />
(Meigen) also occurs under garden compost.<br />
The larvae <strong>of</strong> several species <strong>of</strong> Mydaea develop primarily in fungi: M . affinis Meade<br />
( = discimana Malloch) in Suillus bovinus; M . electa (Zetterstedt) agarics, Boletaceae,<br />
Phallus; M. humeralis Robineau-Desvoidy ( = tincta Zetterstedt) in agarics, boleti,<br />
Morchella; M. maculiventris (Zetterstedt) in Polyporus squamosus; M. orthonevra<br />
(Macquart) ( = detrita Zetterstedt), in Boletus, Suillus, Lactarius, Amanita; M.<br />
setifemur Ringdahl, indet. fungus. M. urbana (Meigen) and M. ancilla (Meigen) have<br />
also been reared from fungi but urbana is found mainly in cow dung and ancilla is also<br />
recorded from human faeces, dog and cattle dung and rotten elm wood. M . corni<br />
(Scopoli) ( = scutellaris Robineau-Desvoidy) breeds in cow dung.<br />
Myospila meditabunda (F.) breeds mainly in cow dung and sometimes in sheep dung;<br />
it has also been recorded from dog dung and human faeces.<br />
Coenosiinae. Larvae <strong>of</strong> Graphomya maculata (Scopoli) (figs 942- 945) were found in<br />
putrid black mud in shallow pools (in France) by Keilin (1917). They were in company<br />
with larvae <strong>of</strong> tipulids, Ptychoptera, tabanids, Eristalis and scathophagids upon which<br />
<strong>the</strong>y were preying. Early authors have recorded larvae <strong>of</strong> this species from dung and<br />
carrion. It has also been found in tree rot-holes (preying on Myathropa, Syrphidae)<br />
and in rotting vegetable matter beside ponds and land drains (Skidmore, 1985). The<br />
larvae have retractile pseudopods on segments 2- 7 and anal papillae.<br />
Most species <strong>of</strong> Spilogona develop in damp soil and moss cushions from which some<br />
have invaded drier situations whilst o<strong>the</strong>rs have become entirely aquatic.<br />
The larva <strong>of</strong> Villeneuvia aestuum (Villeneuve) (figs 946-948) occurs under s<strong>to</strong>nes<br />
(Mercier, 1921) and in wet sand in <strong>the</strong> tidal zone, sometimes in association with and<br />
probably preda<strong>to</strong>ry upon Balanus and possibly o<strong>the</strong>r lit<strong>to</strong>ral animals. The cephalopharyngeal<br />
skele<strong>to</strong>n apparently has <strong>the</strong> pharyngeal sclerite pigmented only anteriorly<br />
(fig. 946).<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Neolimnophora are unknown.<br />
Many <strong>of</strong> <strong>the</strong> larvae <strong>of</strong> Limnophora (figs 949- 950) are aquatic and have caudal<br />
processes which help anchor <strong>the</strong>m <strong>to</strong> dense mosses or liverworts in running water. They<br />
prey mainly on oligochaetes and small insect larvae (Psychodidae, Cera<strong>to</strong>pogonidae,<br />
etc.). L. riparia (Fallen) is common in running water and preys on larval Simuliidae.<br />
Some non-<strong>British</strong> species are known <strong>to</strong> develop in dung or decaying plant and animal<br />
matter.<br />
The larvae <strong>of</strong> all known Lispe species develop in wet sand or mud (including saline<br />
situations) with a high organic content. They are obligate carnivores, probably<br />
dimorphic and usually have <strong>the</strong> posterior spiracles strongly exserted (figs 951- 955).<br />
Vaillant (1953) found that larvae <strong>of</strong> L. consanguinea Loew fed voraciously on tipulid,<br />
chironomid and dolichopodid larvae in <strong>the</strong> labora<strong>to</strong>ry.<br />
Pseudocoenosia abnormis Stein has been reared from wet Sphagnum moss.<br />
The <strong>immature</strong> <strong>stages</strong> <strong>of</strong> Limnospila, Orchisia and Spanochaeta are unknown.<br />
138
Caricea ( = Lispocephala) species have not been reared in Britain, but <strong>the</strong> structure <strong>of</strong><br />
Hawaiian larvae suggests affinities with Spilogona (Skidmore, 1985). <strong>British</strong> species are<br />
likely <strong>to</strong> be aquatic or subaquatic.<br />
Schoenomyza li<strong>to</strong>rella (Fallen) has been reared from puparia 'ex Carex sand' in<br />
Sweden.<br />
Macrorchis meditata (Fallen) has been reared from soil around bulbs imported from<br />
Holland <strong>to</strong> New Zealand.<br />
Larvae <strong>of</strong> Allognota agromyzina (Fallen) were found among wet leaf litter with<br />
numerous tiny oligochaetes on which <strong>the</strong>y were feeding and in company with Uta<br />
sylvatica (Meigen) ( = macroptera (Macquart)) larvae (Tipulidae).<br />
Puparia <strong>of</strong> Dexiopsis lacteipennis (Zetterstedt) have been found amongst seaweed<br />
and those <strong>of</strong> D. minutalis (Zetterstedt) under <strong>the</strong> angiosperm Honckenya peploides,<br />
both on sandy beaches.<br />
Coenosia larvae are obligate carnivores and develop in humus and in plants already<br />
infested with o<strong>the</strong>r Diptera larvae. C. tigrina (F.) is regarded as an important preda<strong>to</strong>r<br />
on Delia antiqua (onion fly) with which it is frequently found in humus soil near plant<br />
roots. The larva <strong>of</strong> C. campestris (Robineau-Desvoidy) ( =sexnotata <strong>of</strong> authors) is<br />
described by Lobanov (1970b), but he gives no biological data. In Austria C. dubiosa<br />
Hennig larvae have been found in noctuid moth larval burrows in stems <strong>of</strong> Typha<br />
latifolia, where <strong>the</strong>y preyed on nema<strong>to</strong>cerous larvae (Sciaridae, Cera<strong>to</strong>pogonidae and<br />
Sca<strong>to</strong>psidae).<br />
S<strong>to</strong>moxyinae. The larvae <strong>of</strong> this subfamily are trimorphic and saprophagous.<br />
S<strong>to</strong>moxys calcitrans (L.) (stable fly or biting house fly) (figs 956--959) develops in<br />
cow-dung. It is ano<strong>the</strong>r important muscid species with a large literature, mainly<br />
because <strong>of</strong> <strong>the</strong> importance <strong>of</strong> <strong>the</strong> blood sucking adult (which attacks man and his<br />
domestic animals) and its disease-carrying potential. The larvae move very rapidly but<br />
feign death if alarmed (Ferrar, 1979).<br />
Haema<strong>to</strong>bia irritans (L.) (horn fly) (adults sit in clusters around <strong>the</strong> horns <strong>of</strong> cattle)<br />
(figs 960--963) and Haema<strong>to</strong>bosca stimulans (Meigen) (cattle biting fly) (figs 964-966)<br />
develop in fresh cow dung and <strong>the</strong> adults suck <strong>the</strong> blood <strong>of</strong> cattle (H. stimulans also<br />
attacks man).<br />
Greenberg ( 1971, 1973) gives a good assessment <strong>of</strong> <strong>the</strong> medical and veterinary<br />
importance <strong>of</strong> <strong>the</strong> S<strong>to</strong>moxyinae, and also <strong>of</strong> o<strong>the</strong>r Muscidae and Diptera in general.<br />
Hippoboscidae<br />
(Figs: larva 967, puparia 1282- 1285)<br />
There are about 200 species <strong>of</strong> Hippoboscidae, which mostly occur in <strong>the</strong> Old World<br />
tropics, but about 35 species are found in <strong>the</strong> western Palaearctic region. Thirteen<br />
species are recorded from Britain, but five <strong>of</strong> <strong>the</strong>se are not resident. Adults <strong>of</strong> about<br />
75% <strong>of</strong> <strong>the</strong> known species occur as ec<strong>to</strong>parasites <strong>of</strong> birds and are commonly known as<br />
'keds'. Hippoboscidae are larviparous, producing a single larva (fig. 967) which<br />
develops internally (3- 8 days), is <strong>the</strong>n released as a prepupa which immediately starts <strong>to</strong><br />
pupariate. The larvae may be extruded on <strong>the</strong> bird, in or around <strong>the</strong> nest or away from<br />
<strong>the</strong> nest. The pupae (figs 1282-1285) are shiny dark brown <strong>to</strong> black, ra<strong>the</strong>r round and<br />
seed-like. Hutson (1984) gives a general account <strong>of</strong> <strong>the</strong> biology, keys <strong>the</strong> adults, and<br />
discusses species likely <strong>to</strong> occur in Britain but not yet recorded. Bequaert (1953-1957)<br />
treats <strong>the</strong> world species.<br />
Ornithomyinae. Adults <strong>of</strong> this subfamily are all ec<strong>to</strong>parasitic on birds.<br />
There is only one <strong>British</strong> record <strong>of</strong> Ornithophila metallica (Schiner) (from <strong>the</strong><br />
whitethroat) and no details <strong>of</strong> <strong>the</strong> life-his<strong>to</strong>ry or <strong>immature</strong>s <strong>stages</strong> are known.<br />
139
Ornithomya species are associated with birds as follows: 0 . avicularia (L.) (fig.<br />
1282) on larger passerines (e.g. thrush, starling size upwards), Ciconiiformes,<br />
<strong>An</strong>seriformes, Accipitriformes, Falconiformes, Galliformes, Gruiformes, Charadriformes,<br />
Columbiformes, Cuculiformes, Strigiformes, Caprimulgiformes, Piciformes;<br />
0. fringillina Curtis on small Passeriformes up <strong>to</strong> size <strong>of</strong> greenfinch particularly<br />
Motacillidae, Prunellidae, Turdidae, Muscicapidae, Fringillinidae, Embezeridae; 0.<br />
chloropus Bergroth (lagopodis Sharp) (<strong>the</strong> grouse fly) (fig. 1283) on birds <strong>of</strong> open<br />
habitat particularly Accipitriformes, Falconiformes, Galliformes, Charadriformes,<br />
Cuculiformes, some Passeriformes; 0 . biloba Dufour is on swallows. Hill has studied<br />
<strong>the</strong> pupae ( 1962) and life-his<strong>to</strong>ries ( 1963) <strong>of</strong> Ornithomya.<br />
Crataerina species are associated with birds as follows: C. hirundinis (L.) (house<br />
martin, occasionally sand-martin, swallow and swift), C. pallida (Latreille) (swift,<br />
occasionally hirundines or o<strong>the</strong>r passerines). Overwintering pupae may be found in <strong>the</strong><br />
nests while <strong>the</strong> birds are away from Britain.<br />
!costa is <strong>the</strong> largest genus <strong>of</strong> <strong>the</strong> family (56 species) <strong>of</strong> which only two occur (rarely) in<br />
Britain associated with birds as follows: /. ardeae (Macquart) on Ardeidae (in Britain<br />
on purple heron, bittern and little bittern); /. minor Bigot (tree pipit and corncrake,<br />
both on Fair Isle).<br />
0/fersia spinifera (Leach) (fig. 1284) is known from only one specimen recorded in<br />
Britain on a frigate bird. Guimaraes (1944) describes <strong>the</strong> larva and Bequaert (1957)<br />
figures <strong>the</strong> puparium.<br />
Hippoboscinae. Adults <strong>of</strong> this subfamily are ec<strong>to</strong>parasites <strong>of</strong> mammals.<br />
Hippobosca equina L. (forest fly) is on horses and occasionally cattle, man and dogs.<br />
Lipopteninae. Lipoptena cervi (L.) (deer fly) (fig. 1285) occurs principally on roe deer,<br />
and red deer; also on fallow deer, sika deer and reindeer with stray records from badger,<br />
man and dog.<br />
Melophagus ovinus (L.) (known variously as skeep-ked, sheep-fly, sheep-louse,<br />
sheep-tick) occurs on sheep where <strong>the</strong> larva sticks <strong>to</strong> <strong>the</strong> fleece <strong>of</strong> <strong>the</strong> host and <strong>the</strong> whole<br />
life his<strong>to</strong>ry is passed <strong>the</strong>re.<br />
N ycteribiidae<br />
(Fig: puparium 1286)<br />
There are over 250 species <strong>of</strong> Nycteribiidae (bat-flies), most <strong>of</strong> which are found in<br />
<strong>the</strong> Old World, especially in <strong>the</strong> Oriental and Pacific areas. About 12 species occur<br />
in Europe <strong>of</strong> which three have been recorded in Britain. Females leave <strong>the</strong> host bat<br />
<strong>to</strong> deposit <strong>the</strong> fully grown larva (as a non-feeding prepupa) on beams, walls, etc.<br />
near <strong>the</strong> bat colony. The larva is hemiovoid. Sticky secretions, helped by a narrow<br />
marginal 'skirt', secure <strong>the</strong> flat ventral surface firmly <strong>to</strong> <strong>the</strong> substrate. In addition,<br />
peristalsis <strong>of</strong> <strong>the</strong> freshly deposited larva ensures an airtight seal in which <strong>the</strong> female<br />
assists by backing over it and pressing down with her body. The freshly deposited<br />
larva is white and s<strong>of</strong>t but soon hardens <strong>to</strong> a dark reddish-brown puparium (fig.<br />
1286). Hutson (1984) gives a general account <strong>of</strong> <strong>the</strong> biology and keys <strong>the</strong> adults.<br />
Immature <strong>stages</strong> are described by H . Scott (1934), Ryberg (1939) and Schultz<br />
(1938).<br />
Nycteribia kolenatii Theodor & Moscona is mostly restricted <strong>to</strong> Dauben<strong>to</strong>n's bat<br />
(Myotis dauben<strong>to</strong>ni (Kuhl).<br />
Phthiridium biarticulata (Hermann) occurs mainly on horse-shoe bats (Rhinolophus<br />
spp.) but also occur on long-eared bats (Plecotus spp.).<br />
Basi!ia nana Theodor & Moscona is found on <strong>the</strong> rare Bechstein's bat (Myotis<br />
bechsteini Kuhl)).<br />
140
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