Royal Entomological Society HANDBOOKS FOR THE IDENTIFICATION OF BRITISH INSECTS To purchase current handbooks and to download out-of-print parts visit: http://www.royensoc.co.uk/publications/index.htm This work is licensed under a Creative Commons Attribution-NonCommercial-ShareAlike 2.0 UK: England & Wales License. Copyright © Royal Entomological Society 2012 Handbooks for the Identification of British Insects Vol. 7, Part 11 CLASSIFICATION AND BIOLOGY OF BRACONID WASPS (HYMENOPTERA: BRACONIDAE) M. R. Shaw and T. Huddleston ROYAL ENTOMOLOGICAL SOCIETY OF LONDON flandbooksforthe Identification of British Insects Vol. 7, Part 11 Editors: W. R. Dolling & R. R. Askew CLASSIFICATION AND BIOLOGY OF BRACONID WASPS (HYMENOPTERA: BRACONIDAE) By M. R. Shaw Department of Natural History, Royal Museum of Scotland, Edinburgh EHllJF. and T. Huddleston / Department of Entomology, Natural History Museum, London SW7 5BD. 1991 ROYAL ENTOMOLOGICAL SOCIETY OF LONDON The aim of the Handbooks is to provide illustrated identification keys to the insects of Britain, together with concise morphological, biological and distributional information. The series also includes a Check List of British Insects. Each part should serve both as an introduction to a particular group of insects and as an identification manual. Details of handbooks currently available can be obtained from Publications Sales, The Natural History Museum, Cromwell Road, London SW7 5BD. World List abbreviation: Handbk !dent. Br. Insects ©Royal Entomological Society of London, 1991 First published 1991 by the British Museum (Natural History) at TheN atural History Museum, London ISBN 0 901546 77 1 Photoset by Row land Phototypesetting Ltd Bury St Edmunds, Suffolk Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset Contents Page Introduction . 3 Additional literature . 6 General biology . 7 Phylogeny and classification. 16 Morphological terminology . 19 Recognition ofBraconidae . 24 Checklist ofthe British genera . 27 Key to subfamilies of British Braconidae . 30 Accounts of British subfamilies . 44 Adeliinae . 44 Agathidinae . 44 Alysiinae . 46 Aphidiinae . 52 Blacinae . 54 Braconinae . 54 ( Cardiochilinae) . 58 Cenocoeliinae . 58 Cheloninae . 59 Doryctinae ....................................... 62 Euphorinae ..................................... 65 Gnamptodontinae ................................. 69 Helconinae ....................................... 70 Histeromerinae ................................... 72 Homolobinae ..................................... 73 Ichneutinae ..................................... 74 Macrocentrinae ................................... 74 Meteorinae ....................................... 77 Microgastrinae ................................... 80 Miracinae ....................................... 87 Neoneurinae . 87 Opiinae . 87 Orgilinae . 91 Rogadinae . 91 Sigalphinae . 96 Acknowledgements . 97 References . 97 Glossary . 125 Introduction Hymenoptera constitute one of the largest and most successful orders of insects. The order is divided into two sub-orders, Symphyta ( sawfiies) and Apocrita. The larger of these is Apocrita, which is best known to the layman for the remarkable levels of social organisation attained by some species of bees, wasps and ants, the so-called aculeate Hymenoptera. However, apocritans are of at least equally great significance and interest on account of the parasitoid lineages that have evolved as the so-called parasitic Hymenoptera. In fact, these two groups are rather arbitrary divisions of the 3 sub-order Apocrita, and have much in common with one another biologically, as is stressed by Gauld & Bolton (1988) in their comprehensive introductory account of the biology and systematics of the order Hymenoptera. Typically the parasitoids, the 'parasitic Hymenoptera', develop as larvae by feeding on the immature stages of other insects: the fact that the latter are killed in the process is one of several features that set parasitoids (known also as protelean parasites) apart from conventional parasites (Askew, 1971). In parasitoids the adult stage is free-living, though it has the job of locating the 'host' for the next generation. The larvae of parasitoids differ from typical predators in that they do not pursue successive prey items, but are generally rather helpless creatures dependent on their mother to place them, as eggs, on or in the body of a single 'host', which must then provide sufficient food for all their growth to take place. There are several rather minor variations on this life-style (e.g. attacking adult instead of immature insects; consuming spiders instead of insects; feeding on successive host eggs in a concealed batch, thereby inflicting more than the usual single host mortality; or having first-instar larvae or even eggs that make first contact with the host), and the biological distinction between the 'parasitic' and the 'aculeate' Hymenoptera can run very thin. This is chiefly because all apocritans share and still exhibit the ancestry of a free-living adult form that provides for a larva feeding on a high-protein diet that it cannot locate for itself (Gauld & Bolton, 1988). Indeed, some species in 'aculeate' groups behave as typical parasitoids in developing on a single 'prey' item which, in extreme cases, may have been scarcely or not at all moved by the female parent and might even recover its activity for a while. In Britain there are about 6500 species of Hymenoptera- more than in any other insect order and just one of the parasitoid superfamilies, Ichneumonoidea, contains nearly half of them. Most workers now recognise only two families in Ichneumonoidea. These are Ichneumonidae, with over two thousand British species, and Braconidae, with nearly 1200 representatives in Britain. They are, respectively, the largest and the second largest families of British insects, with only Staphylinidae (Coleoptera) otherwise near the thousand mark. Ichneumonoids also include the biggest parasitoids. Although in size British Ichneumonidae (mostly 6-17 mm long) tend to over­ shadow Braconidae (mostly2-6 mm in Britain), both groups contain some large and attractive species and Braconidae exhibit the greater range of morphological diver­ sity. All braconids are parasitoids of other insects, developing as larvae on the tissues of the host, which is killed as a result of the association. They are mostly very host-specialised, being adapted to attack and feed as larvae on a narrow range of hosts that is delimited according to a complex and fascinating mix of extrinsic and intrinsic factors. In attaining this intimate association with their hosts, they have evolved specialised and often remarkable behavioural and other biological adapta­ tions, manifest to a greater or lesser extent in the adult's morphology. They are important in the population dynamics of their hosts, many of which are plant-feeding insects, and braconids have been particularly heavily involved in the so-called classical biological control of insect pests (and more recently in integrated pest management programmes), often with considerable success (Clausen, 1978). Being so specialised also gives braconids, along with all parasitoids, a high potential as indicators of environmental richness and stability. In view of their numbers, interesting biology and economic significance it is surprising that Braconidae (and indeed Ichneumonidae) are studied by so few entomologists. Even in Britain, where knowledge of our insect fauna in general is unrivalled, very little is known about the biology or distribution of the great majority of the species belonging to the two largest families. Why has the enthusiasm of amateur entomologists, on whose painstaking fieldwork so much of our knowledge of British insects has depended, so rarely been channelled into studying the larger 4 parasitic Hymenoptera? It is certainly not because they are uncommon or unimpor­ tant; but it may be because of a lack of introductory texts. Without an introduction, they have perhaps seemed hopelessly difficult and obscure even to the point of dullness. Several recent publications, particularly Gauld & Bolton (1988), have made Hymenoptera more accessible to non-specialists. From that base, the purpose of this Handbook is to bring Braconidae into focus: to provide access to the information needed to study them in Britain confidently and to best effect. Rather than take up space on topics that have been fully discussed elsewhere, such as ways of collecting, rearing, mounting and storing specimens (e.g. Gauld & Bolton, 1988; Fitton et al., 1988; Shaw, 1990) and the general biology of parasitoids (Askew, 1971) or Hyme­ noptera (Gauld & Bolton, 1988), we have concentrated on two main aims. First, we have tried to provide a simple and workable illustrated key to the 24 subfamilies of Braconidae known to occur in Britain, and a guide to the literature that is available for species-level identification for each of these. We hope that this will make trying to identify braconids much easier. Second, we have attempted to summarise the host-associations and biology of each subfamily at the level of current knowledge, as far as possible pointing out what important aspects are not known as well as summarising those that are. We hope that this account, which occupies the greater part of this work, will encourage interest in the biology of Braconidae. In drawing together a lot of otherwise scattered literature we also hope to provide an easy means for the user to relate new information,