Plant Protection Quarterly Vol.16(3) 2001 129 the control of Cassinia spp. was first re- ported by Campbell and Wykes (1991, Biological control of weedy native plants in Australia 1992). Campbell et al. (1994) and Holt- kamp and Campbell (1995) reported on Eligio Bruzzese and Ian Faithfull, Keith Turnbull Research Institute, the impact of a redistribution program of Department of Natural Resources and Environment, PO Box 48, Frankston, Austrotachardia sp. and Paratachardinia sp. Victoria 3199, Australia. scales and on rudimentary host specificity tests which indicated that Austrotachardia sp. was specific to C. arcuata, C. quinque- faria R.Br. and C. longifolia. Campbell et al. Abstract (1994) also commented on other native in- Biological control of weeds is the use of without the natural enemies that suppress sects which they observed attacking selected natural enemies of a plant to them in their area of origin. There is how- Cassinia spp. suppress its population to a more accept- ever a widespread belief that biological More recently, Sparks and Robinson able level, in areas where the plant is un- control is not possible or is inappropriate (1997) investigated over 40 arthropods for desirable. It is a cost-effective weed man- for native organisms (Carl 1982). Only a biological control of weedy rangeland agement technique for larger infesta- handful of plants that are considered na- shrubs in western New South Wales and tions of weeds that have a lower priority tive to a country have been the targets of south west Queensland and found two for control by more rapid techniques biological control programs: Julien and with potential, the soft scale, Pulvinaria sp. such as herbicides. In Australia, as in Griffiths (1998) list 23 weeds worldwide (Coccidae) for Eremophila mitchellii Benth. other countries, the main thrust of bio- that have been targeted in a country where and Eremophila deserti (A.Cunn. ex Benth.) logical control utilizes the classical ap- the plant is in part of its native distribu- Chinnock (Myoporaceae) and the hop- proach that targets invasive exotic plants tion. Only two of these are in Australia. bush mite (an unnamed eriophyid) for introduced without the natural enemies Unfortunately in the last 200 years the hopbush, Dodonea viscosa ssp. angustissima that suppress them in their area of origin. intentional movement of native plants for (DC.) J.G.West (Sapindaceae). Sparks There is, however, a widespread belief horticulture, forestry and urban landscap- (2000) reported that both agents estab- that biological control is not possible or ing, coupled with changes in ecosystem lished at all release sites. In the short 18 is inappropriate for native plants. Conse- management, have assisted a number of month study period, Pulvinaria sp. did not quently only a small number of plants native Australian plants to become inva- have an impact on the target plants while considered native to a country have ever sive within Australia. An important factor the mite killed up to 17% of hopbush in been the targets of biological control and preventing plants becoming weeds in the study plots and was considered to examples in Australia are rare. their native habitat is the presence of bio- have great potential. In the last 200 years the intentional control agents that have evolved with the In all these cases the biological control movement of native plants for horticul- plant. Once released, or partially released works involved the redistribution and ture, forestry and urban landscaping, from predators and parasites through the augmentation of natural enemies that coupled with changes in ecosystem man- actions of man, some native plants may were already present in the general area agement, have assisted a number of na- obtain an advantage over the local flora but were too slow at colonizing stands of tive Australian plants to become damag- and become invasive weeds. the unwanted shrubs. The plants were in- ingly invasive within Australia. This pa- This paper looks at past and current ef- vasive because of changed grazing and per looks at current efforts on biological forts to biologically control weedy native fire management regimes. control of weedy native plants in Aus- plants in Australia and discusses possibili- tralia and discusses possibilities for tar- ties for targeting additional native plants Possible targets geting additional species that are consid- that are considered weedy. Current proto- Which native invasive plants could be tar- ered weedy. Current protocols for bio- cols for biological control of exotic weeds geted for biocontrol? It is proposed that logical control of exotic weeds are exam- and their use in biological control of na- three main types of native weeds occur in ined and their use in biological control tive plants are examined and other insti- Australia: of native plants is discussed. We seek to tutional and legal constraints in which na- • Plants that increase in density in their demonstrate in principle that safe tive plant biocontrol may be regulated are current area of distribution due to biocontrol of native plants in Australia is discussed. We seek to demonstrate in changed management regimes. technically feasible and propose an insti- principle that safe biocontrol of native • Plants that extend their current geo- tutional and legal framework in which it plants in Australia is technically feasible graphic range due to changed manage- can be regulated. and to define the framework in which it ment regimes. These lend themselves may be regulated. to redistribution of native agents. Introduction • Plants that have escaped their natural Biological control of weeds is the use of Past and present research in enemies because they are now sepa- selected natural enemies of a plant to sup- Australia rated from their natural area of distri- press its population in areas where the Very few native plants have been investi- bution by geographical barriers. These plant is undesirable. It is a cost-effective gated for biological control in Australia. lend themselves to classical biocontrol. weed management technique for larger We have been able to find two examples infestations of weeds that have a lower in the literature totalling five species of Invaders of disturbed areas priority for control by more rapid tech- plants. Julien and Griffiths (1998) list Chi- Terrestrial plants that increase in density niques such as herbicides. A large number nese scrub or Sifton bush (Cassinia arcuata in their current area of distribution due to of invasive plants around the world are R.Br.) and coughbush (Cassinia laevis changed management regimes are usually currently targeted by biological control R.Br.) (Asteraceae) in which species of unpalatable species which are able to in- programs (Julien and Griffiths 1998). In native scale insects (Kerridae or Laccifer- crease quickly and invade disturbed habi- Australia, as in other countries, the main idae) were redistributed to areas where tats resulting from land clearing, the intro- thrust of biological control utilizes the these native shrubs were invasive in ex- duction of grazing animals and changed classical approach that targets invasive tensive pasture situations. The possibility fire regimes. They are usually rangeland exotic plants that have been introduced of redistributing native scale insects for shrubs unpalatable to sheep or cattle and 130 Plant Protection Quarterly Vol.16(3) 2001 are considered weeds by the extensive by taking advantage of anthropogenic areas of south-west Western Australia and grazing industries. Two scenarios occur changes that allow them to grow in areas south-eastern Australia. The differences with such species. Either they are able to previously unsuitable for them. The new between these regions are of the same or- escape their full suite of natural enemies areas may be less climatically or ecologi- der as those between Europe and North as they increase in density, or, the distri- cally suitable to some of their natural en- America (Doing 1981). Many examples bution of their natural enemies extends emies or their natural enemies are actively occur of eastern Australian plants that are with them but the plant remains at a den- or unintentionally suppressed as part of invasive in Western Australia and vice sity that is considered ‘weedy’ because it the changed management regime. Pitto- versa. These plants have often been delib- takes the place of more palatable species sporum undulatum Vent. in eastern Aus- erately introduced and behave in the same utilized by domestic animals. Data that tralia is one of the best known examples manner as exotic weeds. Coast tea tree, verify these scenarios is largely unre- (Gleadow 1982) but again, comparisons of Leptospermum laevigatum (Sol. ex Gaertn.) corded in Australia. The cases of Cassinia the suite of natural enemies of these plants F.Muell. (Myrtaceae), in Western Aus- spp., Eremophila mitchellii, E. deserti and in areas where they are expanding in dis- tralia and bluebell creeper, Sollya hetero- Dodonea viscosa mentioned above indicate tribution and areas where they were found phylla Lindl. (Pittosporaceae), in eastern that some damaging natural enemies such historically have not been carried out. Australia are good examples. as scales and eriophyid mites are unable Such studies are required to determine to readily disperse and increase with the whether redistribution of natural enemies Overseas experience population of the plant and that manual into the expanded range of the plant might The situation of native plants which in- redistribution of some natural enemies provide improved control, but in many vade new bioregions is directly compara- does result in effective suppression of cases these plants are probably suitable ble to the experience with biological con- dense infestations of some native weeds. targets for redistribution of agents. trol of invasive Australian plants in other However, comparisons of the suite of parts of the world, particularly in South natural enemies of these plants in areas Plants in new bioregions Africa (Olckers and Hill 1999) and with where they are considered weedy and in The methods of classical biological control experience in other countries where local areas where they are more in balance with can be applied to plants that have escaped native plants have become weeds (e.g. nature have not been carried out. Why their natural enemies and now have popu- DeLoach 1995). Many Australian native these plants are able to outgrow their lations separated from their natural area plants have become weeds in other parts natural enemies in the short term is not of distribution by geographical barriers of the world and are currently the target known. that their natural enemies cannot cross. of biological control programs (Julien and Australia, being such a large landmass, Griffiths 1998). The information gained by Plants extending their range offers possibilities for classical biocontrol other countries during development of Native plants that rapidly invade new of native plants because some plants that programs for their weeds of Australian habitats and extend their current geo- evolved on one side of the continent are origin may assist us in developing pro- graphic range due to changed manage- now invasive in similar climates on the grams for the same plants when they are ment regimes and have been termed ‘eco- other side. The only major biological bar- invasive in Australia. Table 1 lists some logically out of balance’ (Carr 1993). These rier in mainland Australia is the Nullarbor Australian plants currently the target of plants may be able to escape their full suite Plain and Great Victoria Desert that sepa- biocontrol programs in other parts of the of natural enemies in the longer term rate the southern sub-humid and humid world that are also invasive in Australia.

Table 1. Some Australian plants currently the target of biocontrol programs in other parts of the world that are also invasive in Australia. Species Common name Origin Weed Australia Weed overseas Reference Leptospermum laevigatum Coastal tea tree Vic, NSW, SA Qld, WA, SA S Africa*, NZ Gordon 1999b Acacia dealbata Silver wattle Vic, NSW, Tas SA S Africa*, NZ Dennill et al. 1999 Acacia decurrens Early black wattle NSW Vic, Qld, SA, Tas S Africa*, NZ Henderson 1995 Acacia mearnsii Black wattle Vic, NSW, SA WA S Africa*, NZ, Dennill et al. 1999 Hawaii, Reunion Acacia melanoxylon Blackwood Vic, NSW, Qld, SA, WA S Africa*, NZ Dennill et al. 1999 Tas Acacia longifolia Sallow wattle Vic, NSW, Qld, SA, Vic, SA S Africa*, NZ Donnelly 1995, Tas Dennill et al. 1999 Acacia pycnantha Golden wattle Vic, NSW WA, Tas S Africa* Dennill and Gordon 1991, Dennill et al. 1999 Acacia saligna Orange wattle WA Vic, NSW S Africa* Morris 1997, 1999 Hakea sericea Silky hakea NSW, Qld Vic S Africa*, NZ, Kluge and Neser 1991, Spain Gordon 1999a Hydrilla verticillata Hydrilla Vic, NSW, Qld, NT, south-west WA USA*, Mexico* Balcunias et al. 1995 north-west WA; Europe, Africa, S and E Asia Paraserianthes lophantha Cape wattle WA Vic, NSW, SA, Tas S Africa*, NZ Dennill et al. 1999, Schmidt et al. 1999 Pittosporum undulatum Sweet pittosporum Vic, NSW, Qld Vic, SA, Tas S Africa*, Jamaica, Goodland and Healey St. Helena, Hawaii 1997 * = biocontrol program, NZ = New Zealand. Plant Protection Quarterly Vol.16(3) 2001 131 Potential conflicts of interest Biological Control Act 1984 and corre- candidate agent be also declared an agent Many of the native plants invasive within sponding legislation in each state (e.g. Bio- under the Biological Control Act. Australia have been planted as horticul- logical Control Act 1986 in Victoria), al- tural, forestry or amenity plants. It is lows for the declaration of target species Discussion therefore very likely that if their invasive and of individual biological control Enabling of public consultation is likely to populations are targeted for biological agents. The legislation was developed to increase the costs of such programs but suppression then conflicts of interest will identify and resolve any potential con- will enable a wider range of experts in the arise between members of the community flicts of interest over the targeting of a pest community to comment upon and exam- who consider the plants economically im- for biocontrol and the use of particular ine potential ecological problems and con- portant or desirable in some areas and agents. In addition to the legislation, strict sequences, and will enable better assess- those who consider them to be invasive administrative and scientific protocols are ment of the potential costs and benefits of weeds. Although such conflicts are often currently in place for the declaration of a the programs. Initially we expect that difficult to resolve, we know from over- target weed and the importation, testing community consultation will expose seas experience that for commercially im- and release of exotic biological control widespread resistance to the idea of portant plants it is possible to selectively agents (Paton 1995, Withers et al. 1999). biocontrol of native plants, largely based target the reproductive parts of the plant Nominations of target weeds for bio- on fears of off-target damage. Such resist- with biological control, while preserving logical control are initially presented to ance could be moderated by improving the desirable traits, with minimal impact the Australian Weeds Committee (AWC), our understanding of the extent to which on the commercial values. Such strategies one of the technical committees of the native ‘biocontrol agents’ are becoming aim to reduce the invasive potential of Standing Committee on Agriculture and established around Australia as a result of plants by disruption of plant reproductive Resource Management (SCARM). The unintentional human introductions. The functions. This approach has been suc- nomination is circulated for comment extent of such ‘naturalizations’ is probably cessfully used in South Africa for invasive within each State and if no conflicts of in- much greater than is known. There does species from temperate Australia. For ex- terest are identified then the AWC recom- not appear to be any adequate research ample, some Australian Acacia spp. are of mends that SCARM endorse the weed as a program to systematically document them significant economic importance in South target. If conflicts of interest are identified or to quantify their impact. Opposition by Africa for tan bark and timber, but are also then the recommendation is for the appli- some sections of the community is also extremely serious invasive weeds that de- cant to use the processes of the Biological likely to reduce with a better knowledge grade natural areas and agricultural land, Control Act. Administrative and scientific of the increasing rates at which new exotic and consume vast quantities of water that protocols on the testing and release of ex- organisms that impact on native plants are would otherwise be harvested for domes- otic biological control agents are adminis- being introduced and dispersing. AQIS tic and agricultural use or help restore en- tered by the Australian Quarantine and quarantine interception records and sur- vironmental flows in waterways. Acacia- Inspection Service, in conjunction with veys around major ports provide a sub- feeding insects that reduce flower and Environment Australia, and again involve stantial basis but again the existing knowl- seed production have been deliberately extensive consultation with government edge base appears to be inadequate. In the selected, thus reducing the invasive po- departments in all States. Public consulta- longer term, public consultation should tential of the plants while conserving their tion occurs only if State agencies identify have a positive effect on community atti- desirable attributes (Dennill et al. 1999). conflicts of interest and the procedures of tudes and build up public confidence in Studies are also under way on seed de- the Biological Control Act are brought into the process. stroying agents for Pinus spp. in South operation. In summary we would like to reinforce Africa (Zimmermann and Neser 1999), Because most native plants that are the following points: redistribution of while a feasibility study on Pinus spp. has considered weeds have, or are currently natural enemies of some rangeland shrubs also been carried out in New Zealand being used for horticultural, forestry or has resulted in them being successfully (Brockerhoff and Kay 1998). A similar fea- ornamental purposes, it is highly likely suppressed; additional native weeds sibility study is currently under way on that conflicts of interest will arise if they could be targeted by redistribution of Salix spp. in Australia. (J.L. Sagliocco per- are targeted for biological control. It is natural enemies or classical biocontrol sonal communication). A biological con- therefore proposed that any native plant programs; agents can be selected for spe- trol strategy that seeks to suppress repro- nominated as a target for biological con- cific purposes e.g. target the whole plant duction, but not plant growth itself, would trol should be automatically referred to or more specifically target reproductive be appropriate in Australia to decrease the the Biological Control Act. organs of those species being grown for invasive potential of native forestry spe- Because of the close relationship of na- economic reasons; institutional and legal cies planted outside their natural distribu- tive weeds to non-target species, it is im- frameworks already exist for biocontrol of tion e.g. Eucalyptus globulus Labill. and portant to identify biocontrol agents with exotic weeds and should be used for na- some Acacia and Casuarina species. Bio- very high specificity. We therefore further tive weeds with minor amendment to leg- logical control of Australian native plants propose that protocols for host specificity islation; safe biocontrol of native plants is in Australia can also benefit from reviews testing and releases should follow the cur- technically feasible in Australia and the of overseas experience with Australian rent protocols for exotic agents, developed need for effective biocontrol of such plants phytophagous pests that damage Austral- by AQIS and Environment Australia and is becoming more pressing. Basic studies ian plants in foreign lands. endorsed by experts in government agen- of the natural enemy complexes of major cies from each State and Territory, but native weeds in their natural and intro- Protocols and legislation with an appropriate opportunity for com- duced Australian ranges have not been Existing protocols and legal and institu- munity input. In the case of native weeds undertaken. Such studies should be initi- tional frameworks to regulate biological and native biocontrol agents, there is no ated and we suggest that appropriate control of exotic weeds should be of assist- need for the involvement of AQIS. Envi- candidates would be bluebell creeper (S. ance in resolving conflicts relating to na- ronment Australia should therefore ad- heterophylla) in eastern Australia and tive plant biocontrol. Biological control of minister the protocols and approve the re- coastal tea tree (L. laevigatum) in Western pest species, including weeds, is regulated lease of any agent that is sufficiently spe- Australia. by both Commonwealth and State legisla- cific. Again, because of the likelihood of tion in Australia. The Commonwealth conflicts of interest, it is proposed that the 132 Plant Protection Quarterly Vol.16(3) 2001 Acknowledgments (Entomological Society of Southern Af- Kluge, R.L. and Neser, S. (1991). Biologi- We thank Royce Holtkamp (Agriculture rica). cal control of Hakea sericea (Proteaceae) NSW) for information and assistance with Dennill, G.B. and Gordon, A.J. (1991). in South Africa. Agriculture, Ecosystems obtaining literature, and J.L. Sagliocco Trichilogaster sp. (Hymenoptera: Ptero- and Environment 37, 91-113. (KTRI) for information about biological malidae), a potential biocontrol agent Morris, M.J. (1997). Impact of the gall- control of willows. for the weed Acacia pycnantha (Faba- forming rust fungus Uromycladium ceae). Entomophaga 36, 295-301. tepperianum on the invasive tree Acacia References Doing, H. (1981). Phytogeography of the saligna in South Africa. Biological Control Balcunias, J.K., Purcell, M.F. and Burrows, Australian floristic region. In ‘Austral- 10, 75-82. D.W. (1995). Australian insects as bio- ian vegetation’, ed. R.H. Groves. (Cam- Morris, M.J. (1999). The contribution of the logical control agents for the sub- bridge University Press, Cambridge). gall-forming rust fungus Uromycladium mersed aquatic weed, Hydrilla Donnelly, D. (1995). Host searching be- tepperianum (Sacc.) McAlp. to the bio- verticillata, in the USA. Proceedings of haviour of the seed weevil, Melanterius logical control of Acacia saligna (Labill.) the VIII International Symposium on ventralis: implications for the biological Wendl. (Fabaceae) in South Africa. In Biological Control of Weeds, eds E.S. control of Acacia longifolia in South Af- ‘Biological control of weeds in South Delfosse and R.R. Scott, DSIR/CSIRO, rica. Proceedings of the VIII Interna- Africa (1990–1998), African Entomol- Melbourne, pp. 237-45. tional Symposium on Biological Con- ogy Memoir No. 1’, eds T. Olckers and Brockerhoff, E.G. and Kay, M. (1998). trol of Weeds, eds E.S. Delfosse and M.P. Hill, pp. 125-8. (Entomological So- Prospects and risks of biological control R.R. Scott, DSIR/CSIRO, Melbourne, ciety of Southern Africa). of wilding Pinus contorta in New Zea- p. 577. Olckers, T. and Hill, M.P. (eds) (1999). land. Proceedings 51st New Zealand Gleadow, R.M. (1982). Invasion by Pitto- ‘Biological control of weeds in South Plant Protection Conference, Hamilton, sporum undulatum of the forests of cen- Africa (1990-1998), African Entomol- New Zealand, pp. 216-23. tral Victoria. II Dispersal, germination ogy Memoir No. 1’. (Entomological So- Campbell, M.H. and Wykes, P.J. (1991). and establishment. Australian Journal of ciety of Southern Africa). Possible bio-control of sifton bush by Botany 30, 185-198. Paton, R. (1995). Legislation and its ad- scale insects. Proceedings 6th Confer- Goodland, T. and Healey, J.R. (1997). The ministration in the approval of agents ence Grassland Society of New South control of the Australian tree Pitto- for biological control in Australia. Pro- Wales, Orange, pp. 101-2. sporum undulatum in the Blue Moun- ceedings of the VIII International Sym- Campbell, M.H. and Wykes, P.J. (1992). tains of Jamaica. Research supported by posium on Biological Control of Weeds, Possible bio-control of native weeds by the UK Overseas Development Admin- eds E.S.Delfosse and R.R. Scott, DSIR/ native insects. Proceedings 1st Interna- istration and UK Department of the En- CSIRO, Melbourne, pp. 653-8. tional Weed Control Congress, Mel- vironment Darwin Initiative Pro- Schmidt, F., Hoffmann, J.H. and Donnelly, bourne 2, 106-8. gramme, 24 pp. (Unpublished). D. (1999). Levels of damage caused by Campbell, M.H., Holtkamp, R.H., Gordon, A.J. (1999a). A review of estab- Melanterius servulus Pascoe (Coleo- McCormick, L.H., Wykes, P.J., Donald- lished and new insect agents for the ptera: Curculionidae), a seed-feeding son, J.F., Gullan, P.J. and Gillespie, P.S. biological control of Hakea sericea weevil introduced into South Africa for (1994). Biological control of the native Schrader (Proteaceae) in South Africa. biological control of Paraserianthes shrubs Cassinia spp. using the native In ‘Biological control of weeds in South lophantha (Fabaceae). African Entomol- scale insects Austrotachardia sp. and Africa (1990–1998), African Entomol- ogy 7, 107-12. Paratachardina sp. (Hemiptera: Kerrii- ogy Memoir No. 1’, eds T. Olckers and Sparks, D.R. (2000). ‘Biological control of dae) in New South Wales. Plant Protec- M.P. Hill, pp. 35-43. (Entomological So- woody weeds in western New South tion Quarterly 9, 64-8. ciety of Southern Africa). Wales and within south western Carl, K.P. (1982). Biological control of na- Gordon, A.J. (1999b). Biological control of Queensland’, 19 pp. (Cuttagoa Catch- tive pests by introduced natural en- Australian myrtle, Leptospermum ment Landcare Group, National Land- emies. Biocontrol News and Information 3, laevigatum (J.Gaertn.) F.Muell. (Myrt- care Program, West 2000 Program, 191-200. aceae) in South Africa. In ‘Biological Bourke, NSW, ISBN 0958762546). Carr, G.W. (1993). Exotic flora of Victoria control of weeds in South Africa (1990– Sparks, D.R. and Robinson, N.J. (1997). and its impact on indigenous biota. In 1998), African Entomology Memoir No. ‘Biological control of woody weeds in ‘Flora of Victoria Volume 1. Introduc- 1’, eds T. Olckers and M.P. Hill, pp. western New South Wales and within tion’, eds D.B. Foreman and N.G. 139-43. (Entomological Society of south western Queensland: Final Re- Walsh, pp. 256-97. (Inkata Press, Mel- Southern Africa). port on the NRMS Project No. N4080/ bourne). Henderson, L. (1995). Plant invaders of WN0140.93’. (Cuttagoa Catchment DeLoach, C.J. (1995). Progress and prob- southern Africa. Plant Protection Insti- Landcare Group, Bourke, NSW, ISBN lems in introductory biological control tute Handbook No. 5. ISBN 1-86849- 0958762511). of native weeds in the United States. 026-2. Withers, T.M., Barton Browne, L. and Proceedings of the VIII International Holtkamp, R.H. and Campbell, M.H. Stanley, J. (eds) (1999). ‘Host specificity Symposium on Biological Control of (1995). Biological control of Cassinia testing in Australasia: towards im- Weeds, eds E.S.Delfosse and R.R. Scott, spp. (Asteraceae). Proceedings of the proved assays for biological control’. DSIR/CSIRO, Melbourne, pp. 111-22. Eighth International Symposium on (Department of Natural Resources, Dennill, G.B., Donelly, D., Stewart, K. and Biological Control of Weeds, Canter- Coorparoo, Queensland). Impson, F.A.C. (1999). Insect agents bury, New Zealand, pp. 447-9. (DSIR/ Zimmermann, H.G. and Neser, S. (1999). used for the biological control of Aus- CSIRO Melbourne). Trends and prospects for biological tralian Acacia species and Paraserianthes Julien, M.H. and Griffiths, M.W. (1998). control of weeds in South Africa. In lophantha (Willd.) Nielsen (Fabaceae) in ‘Biological control of weeds. A world ‘Biological control of weeds in South South Africa. In ‘Biological control of catalogue of agents and their target Africa (1990–1998), African Entomol- weeds in South Africa (1990–1998), Af- weeds, 4th edition’. (CABI Publishing. ogy Memoir No. 1’, eds T. Olckers and rican Entomology Memoir No. 1’, eds Wallingford, UK). M.P. Hill, pp. 165-73. (Entomological T. Olckers and M.P. Hill, pp. 45-54. Society of Southern Africa).