Classical Biological Control of Arthropods in Australia

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D.F. Waterhouse

D.P.A. Sands

CSIRo Entomology

Australian Centre for International Agricultural Research

Canberra 2001

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primary mandate is to help identify agricultural problems in developing countries and to commission collaborative research between Australian and developing country researchers in ﬁelds where Australia has special competence.

Where trade names are used this constitutes neither endorsement of nor discrimination against any product by the Centre.

ACIAR MONOGRAPH SERIES

This peer-reviewed series contains the results of original research supported by ACIAR, or material deemed relevant to ACIAR’s research objectives. The series is distributed internationally, with an emphasis on the Third World.

Waterhouse, D.F. and Sands, D.P.A. 2001. Classical biological control of arthropods in Australia. ACIAR Monograph No. 77, 560 pages.

ISBN 0 642 45709 3 (print) ISBN 0 642 45710 7 (electronic)

Published in association with CSIRO Entomology (Canberra) and CSIRO Publishing (Melbourne)

Scientiﬁc editing by Dr Mary Webb, Arawang Editorial, Canberra Design and typesetting by ClarusDesign, Canberra

Printed by Brown Prior Anderson, Melbourne

Cover: An ichneumonid parasitoid Megarhyssa nortoni ovipositing on a larva of sirex wood wasp, Sirex noctilio.

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Foreword

WHEN THE CSIR Division of Economic Entomology, now

Commonwealth Scientiﬁc and Industrial Research Organisation (CSIRO) Entomology, was established in 1928, classical biological control was given as one of its core activities. This was indicative of the emphasis to be placed on biological control in Australia for the foreseeable future and was logical when one considers the potential targets for this approach amongst the many exotic pests of the introduced plants on which Australia still depends almost entirely for its agricultural productivity. Biological control has continued as a mainstay of pest management to the present time, with an impressive number of successes over the years.

The ﬁrst comprehensive review of biological control projects in Australia (which also included those in Papua New Guinea) was that of Wilson (1960). This covered attempts against 53 arthropod pests or groups of pests and 12 weeds. There followed coverage of the world scene by Clausen (1978a), which added brief accounts on Australian projects.

Worldwide projects on weeds have been regularly summarised in an abbreviated form by M.H. Julien (Julien and Grifﬁths 1999) but a comprehensive account of the entire range of arthropod projects in Australia up to the present time, now totalling 98 arthropod pests or groups of pests, has been sorely needed for some time. The authors are to be congratulated on their dedication and persistence in amassing the extensive and scattered information required for the task.

Congratulations are also due to the Australian Centre for

International Agricultural Research (ACIAR), the publisher of this book. This project further extends our close collaboration on biological control activities in the oceanic Paciﬁc and Southeast Asia. ACIAR has already published an impressive number of volumes relevant to the development of signiﬁcant programs (Li Li-ying et al. 1997; Waterhouse 1993a,b, 1994, 1997, 1998; Waterhouse and Norris 1987, 1989; Waterhouse et al. 1999; Klein Koch and Waterhouse 2001; Morris and Waterhouse 2001). One spectacular success has been the effective control of a serious defoliator, the

List of targets

General index

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Australia, by dealing with a threat to Australian agriculture before it reaches our shores.

Classical biological control has the capacity to yield extensive and enduring returns in pest management, though success is not always guaranteed. In their brief overview, the authors estimate an overall success rate of about two-thirds for all projects. This in itself represents a

remarkable return on the scientiﬁc investment made. I warmly commend this volume not only for the wealth of information it contains, but also as an invaluable record of what can and has been achieved by this approach and as an indication of the opportunities that still exist to extend and improve the approach further for Australia’s beneﬁt.

Jim Cullen

Chief, Division of Entomology CSIRO, Canberra

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A Tribute

IN HIS retirement from 1981 to 2000, the late Dr Doug Waterhouse

authored or co-authored 12 books on the biological control, distribution and importance of pests and weeds. These publications are of immense importance and relevance to the objectives of both CSIRO Entomology and ACIAR as they promote the economic, social and environmental beneﬁts to be had with appropriate management of insects. His texts have drawn together relevant information available from as many sources as possible, enabling students and research workers to locate easily, most or all of the information on pests and weeds of Paciﬁc and Southeast Asian countries. The books are essential for planning future biological control projects in the region.

This most recent book by Dr Waterhouse, Classical Biological Control of Arthropods in Australia co-authored with Dr Sands, is the last in the series on regional biological control programs. It covers the history until 1999, of arthropod biological control introductions into Australia, and updates information on biological control projects carried out since the publication by Wilson (1960). Entomologists, including the scientists afﬁliated with CSIRO Entomology, are deeply indebted to Dr Waterhouse for the contributions he has made in all the books published after his retirement. They will be referred to for years to come, guiding new initiatives and recording part of the history of safely and successfully controlling pests and weeds, by classical biological control in Australia and the neighbouring developing nations.

R.J. Clements

Director, ACIAR

List of targets

General index

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AN ACCOUNT is provided of attempts at biological control of arthropod

pests in Australia. Ninety-eight pests or groups of pests have been involved, totalling some 150 species, most of which are exotic. Some 70 were targetted in speciﬁc projects.

The pests are listed alphabetically under Collembola (1), Hemiptera (56), Thysanoptera (1), Orthoptera (2), Coleoptera (9), Diptera (7), Lepidoptera (13), Hymenoptera (4), Acari (4) and Diplopoda (1).

In addition to a summary table of results, a short dossier on each pest species or group provides (a) a precis of the outcomes, together with basic data on biology and pest status, (b) information on native natural enemies and (c) an account of the attempt(s) at biological control and the biology of the most important natural enemies.

Without recent evaluations it is often not possible to assess accurately the level of successful control, but a general overview indicates that about 30 of the target pests are very well controlled and a further 20 are no longer important pests, indicating an overall success rate for target pests of about two-thirds. With the exception of the dung-breeding bush ﬂy, native pests have not proved susceptible to classical biological control.

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Acknowledgments

MANY COLLEAGUES within CSIRO Entomology, State Departments of

Agriculture and universities have provided valuable information and comment, often providing more accurate and up-to-date accounts of the various projects than is available from publications. Although it is not possible to acknowledge all of the many individuals involved, very special thanks are due to Dr M. Carver for much unpublished information on the aphid pests listed. Mr D. Smith of the Queensland Department of Primary Industries (QDPI), Nambour, Queensland provided most valuable information on scales and mealybugs. Mr J. Feehan supplied unpublished information on the distribution and impact of dung beetles and this was supplemented by Drs M. Tyndale-Biscoe, T.J. Ridsdill-Smith and J.N. Mathiessen. Others in CSIRO Entomology include Drs G.H. Baker, J. Daly, P. Greenslade, B.H. Halliday, G.A. Macqueen, W. Milne, R.J. Milner, L.A. Mound, K.R. Norris, J.L. Readshaw, J.P. Spradbery, R.W. Sutherst and K.G. Wardhaugh.

Valuable inputs were also provided by

Queensland G.K. Waite, QDPI Nambour; B.C.G. Scholtz and D.A.H. Murray, QDPI Toowoomba; Prof. R.A.I. Drew, Grifﬁth University;

J.F. Grimshaw and B.M. Waterhouse, Northern Australia Quarantine Strategy (NAQS) Mareeba; K.H. Halfpapp, QDPI, Mareeba; K.J. Chandler and Dr P.G. Allsopp, Bureau of Sugar

Expermiment Stations

New South Wales Dr B.J. Fletcher (retired CSIRO);

Major S.P. Francis, Army Malaria Research Unit; J. McIntosh, Australian Museum

List of targets

General index

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We are most grateful for the valuable data processing inputs provided by Mrs Leanne Slarke and Mrs Bev Johnstone. To Dr Mary Webb, Mrs Kerry Highley and Mr Peter Lynch (ACIAR), we extend our thanks for invaluable editorial advice and help during the later stages of preparation of the text, tables and maps for the book.

It would not have been possible to undertake a major task such as this, when in retirement, without the encouragement, support and patience of Dawn and Susan, our wives, to whom very special appreciation and thanks are due.

Dr T.R. New, La Trobe University South Australia Dr F.D. Morgan, formerly Waite Institute Western Australia A.C. Postle, NAQS, Broome;

G.R. Strickland, Department of Agriculture Northern Territory T.L. Fenner, NT Quarantine;

D. Chin, Department of Agriculture The Natural History Museum,

London

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Introduction

MANY INSECTS that are of little or no economic importance in their

country of origin become important pests when they are introduced to another country without their own natural enemies. Classical biological control is the term used when some or all of these natural enemies are introduced and established in the new country. Areas of the world where much of today’s agriculture is based on introduced crops (e.g. Australia, New Zealand, Hawaii and California) are notable for two facts. One is that a high proportion of their important arthropod pests are exotic; the other is that classical biological control has resulted in many important successes. Worldwide, biological control—either standing alone or as a component of integrated pest management—is attracting increasing interest, partly in order to reduce dependence upon pesticides. Much can be learnt from successes (and failures) with classical biological control projects around the world. A major aim of this publication is to provide ready access, in a single table, to a summary of information on the natural enemies that have been liberated for classical biological control of arthropods in Australia up to 2000, including (when available) their origin, year(s) of liberation, whether they have been established and with what effect. We have adopted, in the ‘Effect’ column of Table 1 page 29, a simple rating system where an introduced natural enemy has become established. The number of ‘+’ symbols in this column is an indication either of (a) when known, the impact of the presence of the natural enemy on the pest population or (b) the resulting abundance of the natural enemy, when the relationship between its abundance and the pest population has not been established. In many cases the indication of effect is a very subjective one, since there has often been no adequate evaluation in recent years, if at all. It is, perhaps, also not surprising that some species have been introduced into quarantine but not liberated. Sometimes this has been due to the very low numbers of living natural enemies imported, to difﬁculties in breeding them on the target host, or a decision not to liberate because of real or apparent lack of speciﬁcity or effectiveness.

List of targets

General index

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microclimates or one group of crops, but far less effective in another set. An example of this is provided by the parasitoid Trissolcus basalis and the once very widespread green vegetable bug, Nezara viridula. The bug is now suppressed by the parasitoid in much of south-eastern New South Wales and Victoria to the extent that it has become an uncommon pest. However, in southern Queensland (particularly where soybeans are grown) and also in a strip of inland New South Wales, extending even into Victoria, it is still regarded as a serious pest. Thus biological control has undoubtedly been highly successful over a vast area, although the pest continues to be a problem elsewhere. It should be pointed out that, whatever the level of suppression brought about by natural enemies, it is highly likely that any contribution they can give to the suppression of pest numbers will be a useful contribution to the integrated management of the pest.

In addition to records in the main table (Table 1 page 29), a brief dossier is provided on each pest and Australian locations mentioned in the text are shown in Map 1 page 22. Only about one-third of the target pests were subjected to major, as contrasted with minor, biological control projects. Since a great deal more information is available on the major attempts than on the remaining targets, these pests are dealt with in considerably greater detail in the individual dossiers. The dossiers outline relevant aspects of biology and pest status and provide details of the attempt(s) at biological control and native natural enemies. It would not be feasible in a book of this size to deal exhaustively with all these aspects. However, key references are provided to facilitate access to the sometimes very extensive literature.

Eleven of the pests have already been dealt with, often in far greater depth, in dossiers published elsewhere (Waterhouse 1993a, 1998; Waterhouse and Norris 1987, 1989) and these have been drawn on extensively in the present accounts of these pests.

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Australia’s ﬁrst involvement with classical biological control was when it served as a source of parasitoids and predators of insect pests (including the Australian cottony cushion scale, Icerya purchasi) that had become established in California and Hawaii. This occurred when A. Koebele visited Australia in 1888 and 1891. The natural enemies sent to California included the predaceous coccinellid beetles

Rodolia cardinalis,Rhyzobius ventralis and Cryptolaemus montrouzieri and the parasitic ﬂies Cryptochetum iceryae and C. monophlebi.

Information on early attempts at classical biological control of insects in Australia was assembled in a valuable review by Wilson (1960), but the names of many of the insects involved have since undergone taxonomic changes. The ﬁrst record of an attempt at establishment of a biological control agent in Australia was of the coccinellid Harmonia conformis sent from New South Wales to Western Australia in 1896. However, it only later became established from Tasmanian adults in 1902, following fruitless liberations in intervening years from New South Wales and Tasmania. Although both South Australia and Tasmania, as well as Western Australia, introduced various coccinellids from other parts of Australia on several occasions before 1900, it was Western Australia that took the lead in relation to the introduction of exotic species when, in 1901, it appointed G. Compere as entomologist to its Department of Agriculture. His task was to introduce natural enemies of the main pests. He travelled widely around the world until 1910, sending back many parasitoids and predators. Wilson (1960) commented that many of these species were not identiﬁed and much of this early work was very poorly documented. Modern classical biological control should certainly not be carried out in this way.

The precise identity of a pest is of minor significance for chemical control, since very few insecticides are sufficiently selective for the exact identity to influence the situation. However, it is quite different for classical biological control. It is then highly desirable (a) that it is the target pest species that suffers by far the greatest adverse effect from the

introduction of a natural enemy, (b) that even close relatives, and especially species less closely related to the pest, are not attacked and (c) if they are attacked, that the impact on their abundance is relatively minor.

List of targets

General index

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found in the arthropod index, referring to its modern counterpart. It is to be expected that further name changes will be made in the future. The name of the author of each valid species is given in the arthropod index.

It is necessary here to issue a word of caution in relation to the validity of the name changes that have been made because, in the majority of cases, it is only the published name of the natural enemy that has been changed. It has not been feasible to check the identity of the natural enemies against voucher specimens and it is possible that the published name has, on occasion, been wrongly applied to the species concerned. Valuable advice on taxonomic problems has been given by colleagues associated with the Australian National Insect Collection, Canberra: Hemiptera, Dr M. Carver; Hymenoptera, J.C. Cardale; Lepidoptera, E.D. Edwards; Diptera, Dr K.R. Norris; Coleoptera, T.A. Weir and Dr E.C. Zimmerman; and Acari, Dr B.H. Halliday.

Taxonomic treatment has generally followed that in Insects of Australia (1991). The present nomenclature has been based, inter alia, on theAustralian Standard List of Common Names (Naumann 1993), the

Catalogue of the Chalcidoidea of the World on CD-ROM (Noyes 1998) and theCABI Arthropod Name Database on CD-ROM (CABI 1995). The nomenclature of mites is based on Halliday (1998) and that of economic plants follows of Lazarides and Hince (1993).

The number of aphid pest species considered to be targets in Table 1

( page 29) cannot be deﬁned easily. This is due to an early adopted practice

of introducing polyphagous parasitoids for a particular target species (e.g.Aphis craccivora), but with the hope that they might also assist in the control of one or more non-target pest species (Carver 1989), by

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With regard to aphid parasitoids (Table 2 page 107):

1. many of the effective parasitoids are polyphagous, although some are restricted to an aphid genus or to a particular habitat (e.g. to aphids on cereals);

2. polyphagous parasitoids have been intentionally introduced that will (or might) attack a range of pest species (Stary 1967b; Carver 1989). Any important pest species attacked by the parasitoid can, therefore, be regarded as a legitimate target of the introduction, and several have been included for this reason; and

3. in earlier years, aphid parasitoids introduced intentionally were often not adequately identified. Whether from this cause, or because they have arrived unaided, the origin and year of arrival of a number of exotic aphid parasitoids that have become established is undocumented. An incomplete picture would be presented if these species were excluded from being listed, so they appear with the symbol (U) for unknown time and method of arrival. At times, additional (often more host-specific) strains of these same species have been introduced to increase the range of pest aphids attacked. In addition, a deliberate attempt was made to establish, on several non-target aphid pest species already present in Australia, parasitoid species that would be established and waiting should the highly damaging Russian wheat aphid, Diuraphis noxia, arrive in Australia (Hughes et al. 1994).

Thus, it was decided to record in Table 1 page 29 a number of pest species which were not primarily or intentionally targetted. These species are annotated throughout the book by the symbol †. Although it would not be reasonable to include such species in calculations of the success of biological control, the information presented is relevant to a consideration of the impact of parasitoids on non-target species.

It was also decided to include attempts to control several native pest species by the introduction of exotic natural enemies, although this procedure does not fit the strict definition of classical biological control. The native species involved are: the Queensland fruit fly, Bactrocera tryoni; the native budworm, Helicoverpa punctigera; several native mosquitoes; the pink sugarcane mealybug, Saccharicoccus sacchari; the wingless grass-hopper, Phaulacridium vittatum; and the black field cricket, Teleogryllus commodus. Another species, the citrophilus mealybug, Pseudococcus calceolariae, is believed to be native to the Sydney region, where it is rare.

List of targets

General index

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arthropods have been included in this book, particularly when the information was incomplete. For example, studies on exotic and native natural enemies of Bemisia tabaci biotype B are current (De Barro et al. 2000), but all are not yet completed or documented (P.J. De Barro, pers. comm.). Other records may have been overlooked. However, we hope that an opportunity will be found to add this additional information to a revised edition of this book.

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List of Targets

COLLEMBOLA 1 Sminthurus viridis 100 HEMIPTERA 2 Acyrthosiphon kondoi 105 3 Acyrthosiphon pisum 114 4 Aleurodicus dispersus 117 5 Aonidiella aurantii 120 6 Aonidiella citrina 120 7 Aonidiella orientalis 126 8 Aphis craccivora 129 9 Aphis gossypii † 132 10 Asterodiaspis variolosa 134 11 Brevicoryne brassicae 137 12 Cavariella aegopodii 139 13 Ceroplastes ceriferus † 141 14 Ceroplastes destructor 144 15 Ceroplastes floridensis † 151 16 Ceroplastes rubens 154 17 Ceroplastes sinensis † 159 18 Chrysomphalus aonidum 163 19 Coccus hesperidum 166 20 Coccus longulus † 170 21 Coccus pseudomagnoliarum † 172 22 Coccus viridis 174 23 Comstockaspis perniciosus 177 24 Diaspis bromeliae † 180 25 Edwardsiana froggatti 182 26 Eriococcus araucariae 184 27 Eriosoma lanigerum 186 28 Hyperomyzus lactucae 189 29 Lepidosaphes beckii † 192 30 Lepidosaphes gloverii † 195 31 Macrosiphum rosae 197 32 Metopolophium dirhodum 199 33 mirid bugs 201

General index

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39 Pineus boerneri 216 40 Pineus pini 216 41 Planococcus citri 219 42 Pseudococcus calceolariae † 226 43 Pseudococcus longispinus 231 44 Pseudococcus viburni 234 45 Pulvinaria polygonata † 236 46 Rhopalosiphum maidis † 238 47 Rhopalosiphum padi † 240 48 Saccharicoccus sacchari 243 49 Saissetia coffeae † 247 50 Saissetia oleae 249

51 Therioaphis trifolii forma clover † 254

52 Therioaphis trifolii forma maculata 256

53 Toxoptera aurantii 260 54 Toxoptera citricidus 260 55 Trialeurodes vaporariorum 263 56 Tuberculatus annulatus 265 57 Unaspis citri 267 THYSANOPTERA 58 Heliothrips haemorrhoidalis 272 ORTHOPTERA 59 Phaulacridium vittatum 274 60 Teleogryllus commodus 277 COLEOPTERA 61 Brontispa longissima 279 62 Bruchus pisorum 282 63 canegrubs 284 64 Cosmopolites sordidus 287 65 Ips grandicollis 290 66 Listroderes difficilis 294 67 Pyrrhalta luteola 296 68 Rhabdoscelus obscurus 298 69 Sitona discoideus 300

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DIPTERA 70 Bactrocera tryoni 305 71 Ceratitis capitata 316 72 Haematobia exigua 322 73 Lucilia cuprina 328 74 mosquitoes 333 75 Musca vetustissima 337 76 Stomoxys calcitrans 348 LEPIDOPTERA

77 armyworms, cutworms and semi-loopers 351

78 Cryptophlebia ombrodelta 357 79 Cydia pomonella 360 80 Grapholita molesta 364 81 Helicoverpa armigera 369 82 Helicoverpa punctigera 369 83 Hellula undalis 377 84 Oncopera spp. 379 85 Phthorimaea operculella 382 86 Phyllocnistis citrella 388 87 Pieris rapae 392 88 Plutella xylostella 399 89 Sitotroga cerealella 405 HYMENOPTERA 90 Caliroa cerasi 407 91 Sirex noctilio 409 92 Vespula germanica 417 93 Vespula vulgaris 417 ACARI 94 Boophilus microplus 420 95 Halotydeus destructor 422 96 Panonychus ulmi 427 97 Tetranychus urticae 429 JULIDA 98 Ommatoiulus moreleti 433

List of targets

General index

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Table 1. Target, including incidental target (†), pests and natural enemies released 29

Table 2. Parasitoids of pest aphids in Australia 107

Table 3. Hyperparasitoids of aphid parasitoids 108

Table 4. Aphid predators recorded in the aphid data base 109

Table 5. Fungi attacking pest aphids in Australia 113

Table 6. Indigenous natural enemies of Aonidiella aurantii and A. citrina 124 Table 7. Indigenous natural enemies of Aonidiella orientalis 128 Table 8. Indigenous natural enemies of Ceroplastes ceriferus 143 Table 9. Indigenous natural enemies of Ceroplastes destructor 150 Table 10. Indigenous natural enemies of Ceroplastes floridensis 153 Table 11. Indigenous natural enemies of Ceroplastes rubens 157 Table 12. Indigenous natural enemies of Ceroplastes sinensis 161 Table 13. Indigenous natural enemies of Chrysomphalus aonidum 165 Table 14. Indigenous natural enemies of Coccus hesperidum 168 Table 15. Indigenous natural enemies of Coccus pseudomagnoliarum 173 Table 16. Indigenous natural enemies of Coccus viridis 176 Table 17. Indigenous natural enemies of Comstockaspis perniciosus 179 Table 18. Indigenous natural enemies of Diaspis bromeliae 181 Table 19. Indigenous natural enemies of Lepidosaphes beckii 194 Table 20. Indigenous natural enemies of Nezara viridula 209 Table 21. Indigenous natural enemies of Parasaissetia nigra 211 Table 22. Indigenous natural enemies of Parthenolecanium persicae 213 Table 23. Hymenopterous parasitoids of Planococcus citri on citrus

at Mundubbera, Queensland 225

Table 24. Hymenopterous parasitoids of Pseudococcus calceolariae 230 Table 25. Hymenopterous parasitoids of Diuraphis noxia in Ukraine,

southern Russia and Georgia 242

Table 26. Natural enemies of Saccharicoccus sacchari 246 Table 27. Indigenous natural enemies of Saissetia coffeae 248 Table 28. Indigenous natural enemies of Saissetia oleae 252 Table 29. Indigenous natural enemies of Unaspis citri 271 Table 30. Indigenous natural enemies of Haematobia exigua 327 Table 31. Natural enemies of Lucilia cuprina and other sheep

and carrion-feeding Calliphoridae 332

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Table 33. Month of first release in each State and the establishment,

where known (*), of exotic dung beetles 346

Table 34. Parasitoids of Stomoxys calcitrans in eastern Australia 350 Table 35. Common Australian cutworms, armyworms and semi-loopers 355 Table 36. Indigenous arthropod natural enemies of armyworms,

cutworms and semi-loopers 355

Table 37. Natural enemies of Cryptophlebia ombrodelta in Australia 358 Table 38. Parasitoids attacking Cydia pomonella in Australia 363 Table 39. Parasitoids attacking Grapholita molesta in Australia 368 Table 40. Indigenous arthropod natural enemies of Helicoverpa armigera and

H.punctigera 373

Table 41. Natural enemies of Pieris rapae 396

Table 42. Natural enemies of Plutella xylostella 404

Table 43. Natural enemies of Halotydeus destructor in southern

New South Wales, 1987–1993 426

Table 44. Some predators of Tetranychus urticae 432

Table 45. Exotic species that have been (or still are) major pests in Australia 439 Table 46. Exotic species that have been (or are still) minor pests in Australia 440 Table 47. Species that are (or may be) native to the Australian region 440

List of targets

General index

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4 Alice Springs 48 Mareeba

5 Alstonville 49 Maryborough

6 Applethorpe 50 Mornington Peninsula

7 Armidale 51 Moe

8 Atherton 52 Moree

9 Bamaga 53 Moruya

10 Batlow 54 Mossman

11 Benalla 55 Mount Isa

12 Bilpin 56 Mundubbera

13 Boigu Island 57 Murray Island 14 Bridgewater 58 Murray River

15 Broome 59 Murray Valley

16 Bundaberg 60 Murrumbidgee Irrigation Area

17 Byfield 61 Murrumbidgee River

18 Cairns 62 Nangiloc

19 Cape York Peninsula 63 Narromine

20 Casino 64 Newcastle

21 Clarence River 65 Norfolk Island

22 Coconut Island 66 Ord

23 Cooktown 67 Palm Island

24 Darnley Island 68 Port Lincoln 25 Dauan Island 69 Prince of Wales Island 26 East Gippsland 70 Queanbeyan 27 Elizabeth Beach 71 Richmond

28 Fremantle 72 Riverina

29 Gippsland 73 Riverland

30 Gosford 74 Rockhampton

31 Goulburn Valley 75 Sabai Island

32 Griffith 76 Sale

33 Grafton 77 Sarina

34 Gympie 78 Seisia

35 Hammond Island 79 Stanthorpe 36 Horn Island 80 Thursday Island 37 Hunter River 81 Torres Strait

38 Innisfail 82 Townsville

39 Kimberley 83 Tully

40 Kuranda 84 Yam Island

41 Lachlan River 85 Yarwun

42 Launceston 86 Yorke Island

43 Lord Howe Island 87 Weipa

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Gre at Di vid in gR an ge (4 7) Gre at Div idi ng Ra ng e(4 7) 3 28 2 15 4 1 68 44 62 5 34 21 33 20 16 49 56 17 74 85 45 77 82 48 38 83 8 46 67 23 55 54 18 9,78 87 35, 36, 69, 80 Torr es Strait (81) 22 13 84,24 57,86 25, 76 7 52 10 43 32 63 12 27 30 71 64 53 70 88 11 76 50 14 42 6,79 Ca pe Yo rk P eninsula (19) 26 29 31 37 Kimber le y (39) Norf olk Island (65) 29 ° 02'S 167 ° 57'E 51 58 41 61 72 59 66

List of targets

General index

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Target arthropod pests

and

natural enemies released

List of targets

General index

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TABLE 1 provides a condensed record of the published attempts at biological control of 98 species or groups of species of Australian arthropod pests. Eight of these species or species groups are native to Australia and 18 have only been incidental targets, leaving 72 species that have been direct targets of classical biological control attempts. Further details are provided in the next section for each target pest.

The pests are arranged alphabetically under each order and the insect orders arranged from the most primitive to the most advanced. More detailed explanations and clariﬁcation of table inclusions are given in the

Introduction.

Explanation of abbreviations and symbols used

: † pest not primarily targetted

? not certain

A blank space indicates that information was either not available or not relevant.

Biological control agent

(U) unknown time and method of arrival; an exotic natural enemy which apparently arrived unaided

(#) introduced primarily against another target pest species (speciﬁed at the end of each pest entry)

Host stage (H.S.) E egg

L larva (or nymph) (with 1, 2 etc. representing larval instar stages—in later tables)

PP pre-pupa P pupa A adult

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Origin of agent/Liberation location N north or northern S south or southern E east or eastern W west or western

ACT Australian Capital Territory NSW New South Wales

NT Northern Territory NZ New Zealand Qld Queensland SA South Australia Tas Tasmania UK United Kingdom USA United States of America Vic Victoria

WA Western Australia Liberation date

Where liberations of a natural enemy have been made at various times spanning a sequence of years (e.g. 1963–72), this does not necessarily indicate that all the intervening years were involved.

Established (Est.)

+ successfully established (+) established, but only brieﬂy

– did not establish

Effect (on pest status in the absence of disruption by pesticides) (Eff.) +++ effective biological control, sometimes only regionally

++ partially effective + little effect – no known effect

List of targets

General index

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Ta

ble 1.

Target, including incidental target (†), pests and na

tural enemies r eleased etc . (f or explana tion of a b br e via

tions and symbols, see pp

. 26–27).

Target Biological control agent

H.S. Origin of agent Liberated Est. Eff. Host(s) References 1. Sminthurus viridus Collembola: Sminthuridae le gume pastur es ACARI BDELLID AE Bdellodes lapidaria (U) L,A Europe < 1931 + + + Womersley 1933 Neomolgus capillatus L,A Netherlands 1965 WA – Wallace 1967, 1974, 1981 France, Morocco 1969 WA + + +

Ireson 1984; Ireson & Paterson 1991

France

1985–90

Tas

+

+++

Ireson & Paterson 1991

ANYSTID AE Anystis wallacei L,A France 1965 WA + + + Waterhouse 1978; Wallace 1981 WA 1976 Vic – Berg 1991 WA 1991 Vic – Berg 1991 WA 1993 Vic +

Gardner & Gardner 1994

WA

1993

Tas

+

Ireson & Webb 1995

2. Ac yr thosiphon k ondoi Hemiptera: Aphididae lucerne, clover HYMENOPTERA APHELINID AE Aphelinus abdominalis (U) L,A Tas 1979 NSW + –

Milne 1986a; M. Carver pers. comm. 1998

BRACONID

AE

Aphidius ervi

L,A

Tas, Japan via USA

1977–79

NSW

+

Milne 1986a; M. Carver pers. comm.

Japan via USA via NZ, Europe

1980–81

NSW

+

+++

Milne 1986a; W.M Milne pers. comm.

Aphidius pisivorus (#) L,A USA 1980 NSW –

M. Carver pers. comm.

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General index

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Ephedrus plagiator

L,A

Japan via USA, NZ

1977–79

NSW

–

Milne 1986a

FUNGI Pandora kondoiensis

(U)

L,A

+

Milner 1982; Milner et al. 1983

Pandora neoaphidis (U) L,A NSW + +

Milner 1978; Milner et al. 1980

(#) intr oduced against Acr ythosiphon pisum 3. Acr ythosiphon pisum Hemiptera: Aphididae lucerne, clover , pea HYMENOPTERA APHELINID AE Aphelinus abdominalis (U) L,A Tas 1979 NSW, Vic + –

Milne 1986a; M. Carver pers. comm. 1998

Aphelinus asychis (#) L,A France 1978 NSW, Vic + – Carver 2000 USA 1980 NSW, Vic – South Africa 1981 NSW, Vic – BRACONID AE Aphidius eadyi L,A via NZ 1980 – Carver 1989 Aphidius ervi (#) L,A

Japan via USA, Europe, Tas

1980–81 NSW, Vic + +++ Milne 1986a, 1999 Aphidius pisivorus L,A USA 1980 NSW, Vic –

Carver 1989; M. Carver pers. comm.

Aphidius smithi

L,A

India via USA

1980–81 NSW, Vic + ? Carver 1989 Aphidius urticae group L,A ? via NZ 1980 NSW, Vic – Carver 2000

FUNGI Pandora neoaphidis

(U) L,A ? ? NSW + + + Milner et al. 1983 (#) intr oduced against Ac yr thosiphon k ondoi Ta ble 1. (cont ’d)

. 26–27).

H.S. Origin of agent Liberated Est. Eff. Host(s) References

(31)

4. Aleurodicus disper sus Hemiptera: Ale yr odidae pol yphag ous HYMENOPTERA APHELINID AE Encarsia sp.

Trinidad via Fiji

1992

Boigu Is.

+

+++

Lambkin 1992, 1996, 1998

Trinidad via Fiji via Torres Strait

1995 Cape York + +++ Lambkin 1998 5. Aonidiella aurantii 6. Aonidiella citrina Hemiptera: Diaspididae citr us, passionfr uit COLEOPTERA COC CINELLID AE unidentified spp. Spain 1903 WA –

Compere 1903; Jenkins 1946; Wilson 1960

unidentified sp. Israel 1904 WA (+) ? Jenkins 1946; Wilson 1960 unidentified sp. China 1905 WA + ? Jenkins 1946 Chilocorus 3 spp. India 1960–63 WA – Houston 1991 Chilocorus bipustulatus L,A Israel 1904 WA + ?

Rosen & DeBach 1978

Chilocorus circumdatus (#) a L,A Hong Kong 1902 WA –

Wilson 1960; Smith & Papacek 1993

India

1960–63

NSW _{via WA}

–

Smith et al. 1997a; Elder et al. 1998; Houston 1991

< 1990 Qld + + + Houston 1991 HYMENOPTERA unidentified sp. China 1902 WA ? Wilson 1960 unidentified sp. Japan 1907 WA ? Jenkins 1946; Wilson 1960 unidentified 2 spp. Sri Lanka 1907 WA – Jenkins 1946; Wilson 1960 Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(32)

unidentified 8 spp. China 1907 WA ? Jenkins 1946; Wilson 1960 unidentified 7 spp. India 1907 WA ? Wilson 1960 unidentified 6 spp. Japan 1909 WA – Jenkins 1946 unidentified >6 spp. Far East 1909 WA ? Wilson 1960 APHELINID AE Aphytis riyadhi USA ?1979 SA – Furness et al 1983; Noyes 1998 Aphytis chrysomphali L China 1905 WA + + +

Wilson 1960; Furness et al. 1983

WA 1925–26 NSW + USA 1954 Vic + Aphytis coheni USA ?1979 SA (+) Furness et al. 1983 Aphytis lingnanensis L China ?1906 WA + +++

WA 1925–26 NSW + USA 1962 Vic – Aphytis lingnanensis (#) b A

Hong Kong via Florida

1977 + + Smith et al. 1995 Japan 1980 + + Thailand 1988 + + Aphytis melinus L,A Pakistan 1961 Vic + +++

Smith 1978a; Furness et al. 1983

Vic 1974 Qld – Coccophagus lycimnia (#) c L,A USA 1907 WA + + Wilson 1960; Noyes 1998 Encarsia citrina L China, USA ca 1900 Vic + + +

Smith et al. 1997a

Israel 1970 + Noyes 1998 Encarsia perniciosi L NSW ca 1900 NSW, Vic, SA + + +

McLaren 1971; Furness et al. 1983; Smith et al 1997a

Israel via USA

1970–73

+

Ta

ble 1. (cont

’d)

. 26–27).

(33)

ENCYR

TID

AE

Comperiella bifasciata (A. citrina

biotype) L,A Japan 1909 WA + +++ Brewer 1971

Comperiella bifasciata (A. aurantii

biotype)

L,A

China via USA

1943–44

all States

+

+++

Jenkins 1946; Wilson 1960; Rosen & DeBach 1978; Furness et al. 1983

1947–49 Qld, Vic, NSW Habrolepsis rouxi L,A USA ?1980 SA, Vic – Furness et al. 1983 EUL OPHID AE Pteroptrix chinensis L China ?1907 ? + + Wilson 1960 1983 NSW (#) intr oduced against aComstockaspis per niciosus ; bUnaspis citri ; cCoccus hesperidum 7. Aonidiella orientalis Hemiptera: Diaspididae pa wpa w COLEOPTERA COC CINELLID AE Chilocorus circumdatus (#) a all Hong Kong 1902 WA –

Wilson 1960; Smith & Papacek 1993

India

1960–63

NSW via WA

–

Smith et al. 1997a

Qld Elder et al. 1998 unknown <1990 + + Houston 1991 HYMENOPTERA APHELINID AE Aphytis chrysomphali (#) b L China 1905 WA + +

Furness et al. 1983; Elder et al. 1998

WA 1925–26 NSW + USA 1954 Vic + Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(34)

Aphytis lingnanensis (#) b L,A China 1906 WA + –

Wilson 1960; Furness et al. 1983; Noyes 1998

WA 1925–26 NSW + USA 1962 Vic + Aphytis melinus (#) b L,A Pakistan 1961 Vic + +

Smith 1978a; Elder et al. 1998; Noyes 1998

Vic 1974 Qld – Encarsia citrina (#) b L China ca 1900 Vic + + + Noyes 1998 Israel 1970 + ENCYR TID AE Comperiella lemniscata A

China, Torres Strait, Qld

1991 Qld + +++ Elder et al. 1998 (#) intr oduced against aComstockaspis per niciosus ; bAonidiella aurantii 8. A phis craccivora Hemiptera: Aphididae le gumes, pol yphag ous HYMENOPTERA APHELINID AE Aphelinus gossypii (U) L,A + + Carver et al. 1993 Aphelinus mariscusae (U) L,A + + Carver 2000 BRACONID AE Aphidius colmani (U) L,A + + +

Grylls 1972; Carver & Stary 1974; Room & Wardaugh 1977; Carver 2000

Aphidius similis

(U)

L,A

+

Carver & Stary 1974

Diaeretiella rapae (U) L,A Europe ? + – Carver 2000 Lysiphlebus fabarum L,A

France, Greece, Turkey

1982–83

+

–

Carver 1984; Carver & Franzmann 2001

Ta

ble 1. (cont

’d)

. 26–27).

(35)

Lysiphlebus testaceipes (U) L,A California 1981–83 + –

Carver 1984; Carver & Franzmann 2001

Trioxys indicus L,A India 1985–86 WA, NSW, Vic – Carver 1989

FUNGI Pandora neoaphidis

(U)

L,A

+

M. Carver pers. comm.

9. A phis gossypii Hemiptera: Aphididae † cotton, cucurbits HYMENOPTERA APHELINID AE Aphelinus gossypii (U) L,A + + + Carver et al. 1993 Aphelinus humilis (U) L,A + – Carver 2000 BRACONID AE Aphidius colemani (U) L,A + +++

Carver & Stary 1974; Room & Wardhaugh 1977

Lysiphlebus fabarum (#) L,A Greece 1982–83 + –

Carver 1984, 1989; Carver & Franzmann 2001

Turkey 1982–83 Lysiphlebus testaceipes (#) L,A California 1981–83 + –

Carver 1984, 1989; Carver & Franzmann 2001

FUNGI Neozygites fresenii

(U)

L,A

<1984

+

Milner & Holdom 1986

(#) intr oduced against A phis craccivora Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(36)

10. Asterodiaspis v ariolosa Hemiptera: Aster olecaniidae oak HYMENOPTERA ENCYR TID AE Habrolepis dalmanni L USA via NZ 1931–32 Tas – Evans 1939b 1933 Tas + + + Wilson 1960

USA via NZ, Tas

1937 ACT, NSW, Vic + + + Evans 1939b 11. Br evicor yne brassicae Hemiptera: Aphididae brassicas COLEOPTERA COC CINELLID AE Adalia bipunctata all 1900s WA + Pope 1988 Coccinella septempunctata all Mediterranean 1903 WA – Wilson 1960 Coccinella undecimpunctata all 1900s WA + Pope 1988 Halmus chalybeus all E. Australia 1902 WA + ? Jenkins 1946 Harmonia conformis all NSW 1896 WA –

Wilson 1960; Clausen 1978a

Tas 1901–02 WA + + Clausen 1978a Orcus lafertei all E. Australia 1902 WA – Jenkins 1946 HYMENOPTERA ? BRACONID AE unidentified 2 spp. a L,A NSW 1902 WA + +++ Jenkins 1946; Wilson 1960 Qld 1902 WA ? ? Wilson 1960 unknown family 1 sp. Sri Lanka 1907 WA + ? Wilson 1960 ‘chaldid’ 1 sp. India 1907 WA + ? Wilson 1960 Ta ble 1. (cont ’d)

. 26–27).

(37)

unknown family 1 sp. ‘Orient’ 1909 WA – Wilson 1960 apr oba bl y Diaer etiella rapae

and possibly the hyper

parasitoid Allo xysta fuscicor nis 12. Ca variella ae gopodii Hemiptera: Aphididae carr ot, umbellifer s HYMENOPTERA BRACONID AE Aphidius salicis L,A California 1962 Vic + +++ Stubbs 1966; Carver 1988, 1989 13. Ceroplastes ceriferus Hemiptera: Coccidae † citr us, per simmon, na ti ve plants HYMENOPTERA APHELINID AE Coccophagus lycimnia (#) a L USA 1907 WA + + Noyes 1998 PTER OMALID AE Scutellista caerulea (#) b E South Africa 1902–03 WA – Jenkins 1946 Sri Lanka ca 1903 WA – Wilson 1960 Brazil 1904 WA – Jenkins 1946 USA 1903–04 WA + +++ Jenkins 1946 Scutellista caerulea (#) c E Uganda, Kenya 1935–38 NSW – Wilson 1960; Sands 1984 (#) intr oduced against aCoccus hesperidum ; bSaissetia oleae ; cCeroplastes destructor Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(38)

14. Ceroplastes destructor Hemiptera: Coccidae citr us LEPIDOPTERA NOCTUID AE Coccidiphaga scitula L,A South Africa 1969–70 NSW – Sands et al. 1986 HYMENOPTERA APHELINID AE Euxanthellus philippiae A N Z 1971 NSW – Sands et al. 1986 ENCYR TID AE Anicetus communis L,A South Africa 1968–72 NSW + + + Snowball 1978 + +++ Sands et al. 1986 1973–74 WA + +

R.G. Lukins pers. comm.

Anicetus nyasicus L,A South Africa 1968–71 NSW + + + Snowball 1978 Qld + +++ Sands et al. 1986 1988 Norfolk Island + +++

D.P.A. Sands unpubl.

Diversinervis elegans A Uganda, Kenya 1935–38 NSW – Wilson 1960 South Africa 1971 NSW + +

Metaphycus helvolus (#) a L USA 1943–47 NSW + + Sands et al. 1986 Qld, SA – Wilson 1960 Microterys nietneri (#) b L,A South Africa 1902 WA + – Prinsloo 1976; Noyes 1998 USA 1907 ? Trichomasthus ingens A South Africa 1969–70 NSW – Sands et al. 1986 Ta ble 1. (cont ’d)

. 26–27).

(39)

EUL OPHID AE Aprostocetus ceroplastae L,A South Africa 1972 NSW + + Sands et al. 1986 PTER OMALID AE

Scutellista caerulea (C. destructor

biotype) A Uganda, Kenya 1935–38 NSW – Wilson 1960 South Africa 1969–70 NSW + + + Sands et al. 1986 (#) intr oduced against aSaissetia oleae ; bCoccus hesperidum 15. Ceroplastes ﬂoridensis Hemiptera: Coccidae † citr us, Ficus spp . HYMENOPTERA APHELINID AE Coccophagus ceroplastae (#) a L,A

Japan via Hawaii

1901

NSW

+

Wilson 1960; D.P.A. Sands unpubl.

Coccophagus lycimnia (#) b L USA 1907 WA + + Noyes 1998 ENCYR TID AE Diversinervis elegans (#) c A Uganda, Kenya 1935–38 ? + +

Sands et al. 1986; Smith et al. 1997a

South Africa 1971 NSW Metaphycus lounsburyi (#) d L,A

South Africa via Israel, Holland

1998 Vic, SA + + Malipatil et al. 2000 Microterys nietneri (#) b A South Africa 1907 WA via USA + + Malipatil et al. 2000 EUL OPHID AE Aprostocetus ceroplastae (#) c L,A South Africa 1972 NSW + + +

Sands et al. 1986; Smith et al. 1997a

PTER OMALID AE Scutellista caerulea (#) d E South Africa 1902–03 WA – Jenkins 1946 Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(40)

Scutellista caerulea (#) d Sri Lanka ca 1903 WA – Wilson 1960 Brazil 1904 WA – Jenkins 1946 USA 1903–04 WA + +++ Jenkins 1946 Scutellista caerulea (#) c Uganda, Kenya 1935–38 WA, NSW ? Wilson 1960 (#) intr oduced against aCeroplastes rubens ; bCoccus hesperidum ; cCeroplastes destructor ; dSaissetia oleae 16. Ceroplastes rubens Hemiptera: Coccidae citr us, mang o, ornamentals HYMENOPTERA CHAL CIDOIDEA unidentified Hawaii 1899 – Wilson 1960 APHELINID AE Coccophagus ceroplastae L,A

Japan via Hawaii

1901

NSW

+

Wilson 1960; Smith et al. 1997a

ENCYR TID AE Anicetus beneficus L,A Japan 1977 Qld + +++ Smith 1986 1990 Norfolk Island + +++

Diversinervis elegans (#) a L,A South Africa 1971 NSW + +

Sands et al. 1986; Loch 1997

Metaphycus helvolus (#) b L USA 1943–47 NSW, Qld, SA + + Wilson 1960; Sands 1984 Microterys nietneri (#) c L,A South Africa 1902 WA + + +

Wilson 1960; Malipatil et al. 2000

USA

1907

Ta

ble 1. (cont

’d)

. 26–27).

(41)

EUL OPHID AE Aprostocetus ceroplastae (#) a L,A South Africa 1972 NSW, Qld + +

Loch 1997; Malipatil et al. 2000

PTER OMALID AE Moranila californica d E NSW 1902 WA + + Perkins 1906 China 1896 Qld – Wilson 1960 Scutellista caerulea (#) b E USA 1904 WA – Wilson 1960 NSW + + + Sands 1984 Qld + + + Smith 1997b (#) intr oduced against aCeroplastes destructor ; bSaissetia oleae ; cCoccus hesperidum dan Australian na ti ve species. R e-intr oduced as Tomocera chinensis 17. Ceroplastes sinensis Hemiptera: Coccidae † citr us, mang o HYMENOPTERA APHELINID AE Coccophagus ceroplastae (#) a L,A

Japan via Hawaii

1901 NSW + + + Wilson 1960; Sands 1984 Coccophagus lycimnia (#) b L USA 1907 WA + + Noyes 1998 ENCYR TID AE Metaphycus helvolus (#) c L USA 1943–47 NSW, Qld, SA + + Snowball 1970; Sands 1984 Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(42)

EUL OPHID AE Aprostocetus ceroplastae (#) d L,A South Africa 1972 NSW, Qld + + Sands 1984 PTER OMALID AE Scutellista caerulea (#) c E South Africa 1902–03 WA – Jenkins 1946 Sri Lanka ca 1903 WA – Wilson 1960 Brazil 1904 WA – Jenkins 1946 USA 1904 WA + +++ Jenkins 1946 Scutellista caerulea (#) d Uganda, Kenya 1935–38 NSW ? Wilson 1960 (#) intr oduced against aCeroplastes rubens ; bCoccus hesperidum ; cSaissetia oleae ; dCeroplastes destructor 18. Chr ysomphalus aonidum Hemiptera: Diaspididae citr

us, palms, ornamentals

HYMENOPTERA APHELINID AE Aphytis chrysomphali (#) L,A China 1905 + + Wilson 1960 Aphytis holoxanthus L,A

Hong Kong via Israel

1974

NSW

+

+++

Snowball & Lukins 1975; Lukins & Snowball 1977a; Smith 1978b

1976 Qld + +++ Encarsia citrina (#) L China ca 1900 + + +

Lukins and Snowball 1977a

Pteroptrix chinensis (U) L China <1907 ? + + Wilson 1960 (#) intr oduced against Aonidiella aurantii Ta ble 1. (cont ’d)

. 26–27).

(43)

19. Coccus hesperidum Hemiptera: Coccidae citr us, passionfr uit, ﬁg , ornamentals HYMENOPTERA unidentified 2 spp. China 1905 WA – Wilson 1960 unidentified spp. China 1906 WA – Wilson 1960 unidentified sp. USA 1907 WA – Wilson 1960 unidentified spp. Sri Lanka 1907 WA – Wilson 1960 unidentified spp. India 1907 WA – Wilson 1960 unidentified 2 spp. Italy 1908 WA – Wilson 1960 unidentified sp. Egypt 1908 WA – Wilson 1960 unidentified spp. China 1909 WA – Jenkins 1946 unidentified spp. India 1909 WA – Jenkins 1946 APHELINID AE Coccophagus ceroplastae (#) a L,A

Japan via Hawaii

1901

NSW

+

+++

India 1907–09 WA Coccophagus lycimnia L USA 1907 WA + + + Wilson 1960; Noyes 1988 Coccophagus semicircularis L ?Europe + +

Smith 1997a; Malipatil et al. 2000

ENCYR TID AE Cristatithorax sp. China 1905–06 WA + + Wilson 1960 Diversinervis elegans (#) b A unknown + +++

Uganda, Kenya 1936–38 NSW + + Encyrtus aurantii ?Europe + + Malipatil et al. 2000 Encyrtus infelix L,A USA + +

Smith 1997a; Malipatil et al. 2000

Ta

ble 1. (cont

’d)

. 26–27).

List of targets

General index

(44)

Metaphycus anneckei (#) c A South Africa 1902 WA + +++ Malipatil et al. 2000 Metaphycus helvolus (#) c L USA 1943–47 NSW, Qld, SA + + +

Metaphycus lounsburyi

(#)

c

L,A

South Africa via Israel & Holland

1998 Qld + + Malipatil et al. 2000 Metaphycus luteolus L USA 1999 Qld + + Malipatil et al. 2000 Microterys nietneri (? = Microterys sp.) L,A South Africa 1902 WA + +

USA 1907 – Malipatil et al. 2000 PTER OMALID AE Scutellista sp. Philippines 1907 WA – Wilson 1960 Scutellista sp. Sri Lanka 1907 WA – Wilson 1960 Scutellista sp. Sri Lanka 1908 WA – Wilson 1960 Scutellista sp. China 1909 WA – Jenkins 1946 Scutellista sp. India 1909 WA – Jenkins 1946 Scutellista caerulea (#) c E South Africa 1902–03 WA – Jenkins 1946 Sri Lanka ca 1903 WA – Wilson 1960 Brazil 1904 WA – Jenkins 1946 USA 1903–04 WA + + Jenkins 1946 Scutellista caerulea (#) b E Uganda, Kenya 1935–38 NSW – Wilson 1960 NSW Malipatil et al. 2000 (#) intr oduced against aCeroplastes rubens ; bCeroplastes destructor ; cSaissetia oleae Ta ble 1. (cont ’d)

. 26–27).

(45)

20. Coccus longulus Hemiptera: Coccidae † citr us, custar d a pple, l ychee, carambola, ﬁg , leucaena, ornamentals HYMENOPTERA APHELINID AE Coccophagus ceroplastae (#) L,A

Japan via Hawaii

1901

NSW

+

(#) intr oduced against Ceroplastes rubens 21. Coccus pseudomagnoliarum Hemiptera: Coccidae † citr us HYMENOPTERA APHELINID AE Coccophagus ceroplastae (#) a L,A

Japan via Hawaii

1901 NSW + + Wilson 1960; Noyes 1998 India 1907–09 WA Coccophagus lycimnia (#) b L USA 1907 WA + +++

Coccophagus semicircularis (#) b L ?Europe + +++

Smith et al. 1997a; Malipatil et al. 2000

ENCYR TID AE Diversinervis elegans (#) c A unknown + + Wilson 1960; Noyes 1998 Uganda, Kenya 1935–38 NSW ? Encyrtus aurantii (#) b ?Europe + + Noyes 1998 Metaphycus helvolus (#) d L USA 1943–47 NSW, Qld, SA + +

Metaphycus luteolus (#) b L USA 1999 Qld + +

Noyes 1998; Malipatil et al. 2000

Ta

ble 1. (cont

’d)

. 26–27).

List of targets

General index

(46)

Metaphycus lounsburyi (#) d L,A South Africa 1902 WA + +

Noyes 1998; Malipatil et al. 2000

Israel 1998 Qld + + Microterys nietneri (#) b L,A South Africa 1902 WA + + Wilson 1960; Noyes 1998 USA 1907 WA – (#) intr oduced against aCeroplastes rubens ; bCoccus hesperidum ; cCeroplastes destructor ; dSaissetia oleae 22. Coccus viridis Hemiptera: Coccidae cof fee, citr us, ornamentals HYMENOPTERA APHELINID AE Coccophagus ceroplastae (#) a L,A

Japan via Hawaii

1901 NSW + + + Malipatil et al. 2000 India 1907–09 WA ? Coccophagus lycimnia (#) b L USA 1907 WA + + Noyes 1998 Coccophagus sp. nr rusti L + +

Smith et al. 1997a

ENCYR TID AE Diversinervis elegans (#) c A Uganda, Kenya 1935–38 NSW + + Malipatil et al. 2000 Diversinervis nr stramineus L,A Kenya 1999 NSW, Qld + +++

Malipatil et al. 2000; D. Smith pers. comm

Encyrtus aurantii (#) b ?Europe + + Noyes 1998 Metaphycus helvolus (#) d L USA 1943–47 NSW + + Wilson 1960; Noyes 1998 Qld, SA – Microterys nietneri (#) b L,A South Africa 1907 WA + +

Wilson 1960; Noyes 1998; Malipatil et al. 2000

USA (#) intr oduced against aCeroplastes rubens ; bCoccus hesperidum ; cCeroplastes destructor ; dSaissetia oleae Ta ble 1. (cont ’d)

. 26–27).

(47)

23. Comstockaspis per niciosus Hemiptera: Diaspididae apple, pear , stone fr uit COLEOPTERA COC CINELLID AE unidentified Spain 1903 WA – Jenkins 1946 Chilocorus circumdatus Hong Kong 1902 WA – Wilson 1960 India 1960–63 WA – Houston 1991 Chilocorus circumdatus (U) <1990 Qld + + +

Houston 1991; D. Smith pers. comm.

Chilocorus stigma 1927 NSW – Wilson 1960 HYMENOPTERA unidentified spp. USA 1907 WA – Wilson 1960 APHELINID AE Aphytis aonidiae + + CSIRO unpubl. Aphytis chrysomphali (#) L China 1905 WA + + +

WA 1925–26 NSW + USA 1954 Vic + Aphytis diaspidis L,A + +

Noyes 1998; CSIRO unpubl.

Encarsia citrina (#) L China ca 1900 Vic + + +

Smith et al. 1997a; Noyes 1998

USA, Israel

1970

+

Encarsia perniciosi (red scale strain) (#)

L China ca 1900 NSW + +

McLaren 1971; Furness et al. 1983

Israel via USA

1970–73 Vic, SA + + Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(48)

Encarsia perniciosi (San José scale strain) Germany 1977 NSW + +

Snowball et al. 1975; Milne & Snowball 1977

Germany 1977–78 Vic + + (#) intr oduced against Aonidiella aurantii 24. Diaspis bromeliae Hemiptera: Diaspididae † pinea pple, br omeliads HYMENOPTERA APHELINID AE Encarsia citrina (#) L,A China ca 1900 Vic + + + Noyes 1998 Israel 1970 + + + Encarsia sp. a + + + Murray 1982a (#) intr oduced against Aonidiella aurantii apossibl y Encar sia citrina 25. Ed wardsiana froggatti Hemiptera: Cicadellidae apple, pr une HYMENOPTERA MYMARID AE Anagrus armatus E USA via NZ 1935 Tas + + + Evans 1937b; Wilson 1960 26. Eriococcus araucariae Hemiptera: Eriococcidae araucaria pines COLEOPTERA COC CINELLID AE Cryptolaemus montrouizieri (#) L,A NSW 1902 WA + +++ Wilson 1960 (#) intr

oduced against Pseudococcidae

Ta

ble 1. (cont

’d)

. 26–27).

(49)

27. Eriosoma lanig erum Hemiptera: Aphididae apple COLEOPTERA COC CINELLID AE Cheilomenes sexmaculata all WA 1911 Vic – Wilson 1960 Exochomus melanocephalus all South Africa 1900 NSW – Wilson 1960 Harmonia conformis all NSW 1896 WA – Jenkins 1946; Wilson 1960 Tas 1901–02 WA + + + Wilson 1960 HYMENOPTERA APHELINID AE Aphelinus mali L,A USA via NZ 1923–25 all States + +++ Wilson 1960 28. Hyperom yzus lactucae Hemiptera: Aphididae lettuce, sowthistle HYMENOPTERA APHELINID AE Aphelinus asychis L,A South Africa 1981 NSW –

Carver & Woolcock 1986

BRACONID AE Aphidius sonchi L,A Japan 1981–83 ACT, NSW –

Carver & Woolcock 1986

France 1981–83 Qld + + +

Carver & Woolcock 1986; CSIRO files

France 1981–82 NSW, Vic, ACT, SA, WA + +++ Sandow 1993 Praon volucre L,A

France, Greece, Turkey

1981–83

ACT,

all States

– + (Tas)

?

Carver & Woolcock 1986; Hughes 1989

Ta

ble 1. (cont

’d)

. 26–27).

List of targets

General index

(50)

FUNGI Pandora neoaphidis (U) L,A unknown unknown + + +

R. Milner pers. comm. 1998

29. Lepidosaphes beckii Hemiptera: Diaspididae † citr us, a vocado , pecans, ornamentals HYMENOPTERA APHELINID AE Aphytis chrysomphali L China 1905 WA + +

Furness et al. 1983; Elder et al 1998; Noyes 1998

WA 1925–26 NSW + + USA 1954 Vic + + Aphytis lepidoaphes L,A China ?1960s ? + +++

Snowball & Sands 1970; Smith et al. 1997a

Aphytis lingnanensis (#) L,A China 1906 WA + +

Lukins & Snowball 1977b; Smith et al. 1997a; Malipatil et al. 2000

WA 1925–26 NSW + ? USA 1962 Vic + ? Encarsia citrina (#) L China ca 1900 Vic + + +

Snowball 1967; Smith et al. 1997a

Israel 1970 + + EUL OPHID AE Pteroptrix chinensis L China ?1907 ? + + Wilson 1960 (#) intr oduced against Aonidiella aurantii Ta ble 1. (cont ’d)

. 26–27).

(51)

30. Lepidosaphes glo verii Hemiptera: Diaspididae † citr

us, macadamia, cocon

ut, palms COLEOPTERA COC CINELLID AE Chilocorus circumdatus (#) a L,A Hong Kong 1902 WA –

Wilson 1960; Smith & Papacek 1993; Smith et al. 1997a; Elder et al. 1998

India 1960–63 WA – <1990 NSW via WA + + Qld + + Houston 1991 HYMENOPTERA APHELINID AE Aphytis chrysomphali (#) b L China 1905 WA + + +

Wilson 1960; Furness et al. 1983; Noyes 1998

WA 1925–26 NSW + + USA 1954 Vic + + Aphytis lingnanensis (#) b L,A China ?1906 WA + +++

Aphytis lingnanensis (#) c WA 1925–26 NSW + + USA 1962 Vic –

Hong Kong via Florida

1977–78

Qld

–

Smith et al. 1995; Noyes 1998

Japan 1981–82 Qld + + Field 1984 Thailand 1988–90 Qld + +

Smith & Papacek 1993

(#) intr oduced against aComstockapsis per niciosus ; bAonidiella aurantii ; cUnaspis citri Ta ble 1. (cont ’d)

. 26–27).

List of targets

General index

(52)

31. Macrosiphum rosae Hemiptera: Aphididae roses DIPTERA SYRPHID AE unidentified L,A Philippines 1907 WA – Jenkins 1946 HYMENOPTERA APHELINID AE Aphelinus gossypii (U) L,A + – Maelzer 1977 BRACONID AE Aphidius ervi (#) L,A

Tas, Japan via USA; Japan via USA via NZ, Europe

1908–81 SA + + Milne 1991 Aphidius rosae L,A Italy 1993 SA + + +

Kitt & Keller 1998

(#) intr oduced against Ac yr thosiphon k ondoi 32. Metopolophium dirhodum Hemiptera: Aphididae roses, cer eals HYMENOPTERA BRACONID AE Aphidius ervi (#) L,A

Tas, France, Greece, Italy, California, NZ

1977–81 + + Carver 2000 Aphidius rhopalosiphi L,A

France via Chile

1985–86

ACT, NSW, Vic

–

CSIRO files

UK & France via NZ

1987–88 ACT, NSW, Vic, Tas + +++ CSIRO files (#) intr oduced against Ac yr thosiphon k ondoi Ta ble 1. (cont ’d)

. 26–27).

(53)

33. mirid bugs Hemiptera: Miridae

cotton

FUNGI Beauveria bassiana

L,A

USA

1994

NSW

–

AQIS 1999b; R.K. Mensah pers. comm.

34. Myzus per sicae Hemiptera: Aphididae † peach, ve geta bles etc . HYMENOPTERA BRACONID AE Aphidius colmani (U) L + + Carver 1989, 1995 Aphidius similis (U) L + +

Stary & Carver 1979

Ephedrus persicae (U) L + + Carver 1989 35. Ne zara viridula Hemiptera: P enta tomidae maize, cotton, le gumes, sorghum, grains, ve geta bles, n uts DIPTERA TACHINID AE Bogosia antinorii L,A Kenya 1958 – Greathead 1971 Trichopoda giacomellii L,A Argentina 1996–99 + +++

Coombs & Sands 2000

Trichopoda pennipes L,A Florida 1941–43 – Wilson 1960 1949–50 – 1952–53 – 1980 – Michael 1981 Hawaii 1980 – Ta ble 1. (cont ’d)

. 26–27).

Classical Biological Control of Arthropods in Australia

D.F. Waterhouse

D.P.A. Sands

Foreword

List of targets

General index

Contents

A Tribute

List of targets

General index

Acknowledgments

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General index

Introduction

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List of Targets

General index

List of targets

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List of targets

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List of targets

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Target arthropod pests

and

natural enemies released

List of targets

General index

Explanation of abbreviations and symbols used

List of targets

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List of targets

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List of targets

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List of targets

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List of targets

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List of targets

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List of targets

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List of targets

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List of targets

General index

List of targets

General index