Biological control of weeds and the dried fruits industry 05-3 pp091-97 Delfosse.pdf · 14....

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Biological control of weeds and the dried fruits industry Ernest S. Delfosse, CS IRO Division of Entomology, GPO Box 1700, Ca nb erra, ACf 2601, Australia . Summary I propose that weed species of importance to the dried fruits industry can be divided into three groups. The first group co ntains the most important weeds to the industry: Emex spp. three- co rnered jack, E. aus- tralis); puncturevine or caltrop (Tribulus ler- restTis ); and gentle Annie lind related Cenchms spp. The second group co ntains weeds of lesser importnnce to the industry, hut because of their importance to other sec: t ors of Australian agriculture and the environment, ure cu rrent targets for bio- logical control programs: skeleton weed (ChonLirilia jllncea), Bathurst burr (Xant" - ium sp;nosllIn )j Noogoonl burr (X. s lrumar;um )j thistles (CarduIlS, Cirsium, Onopordum and Silyhllln spp.)j 1)lIterson's curse (EcI,;um plan/ag;newn); com mon heliotrope (Helio/ropiutn eur opaewn)j St. John's wort (Hypericum perJoraJmn )j docks and sorrels (Rum ex spp.) j and silverleaf nightshade (Solan um elaeagnifoliwn ). The third group contains a large number of weeds of oct..'lsional, sporadic or geographi- cally-limited importan ce to the industry, some of which are turgets for biological con- trol programs in early stuges of develop- ment. Discli ssion of biological control pro- grams provides inf or mation on the types of weeds and biologicnl control Hgents cur- rently studied by CS IRO. Also discussed briefly are uspects of foreign exploration, opportunities for co-operation between Ru- ral Industry Research Funds, and co-opera- tion between CSIRO and the Stat es in im- plementHtion of the programs. Introduction Thcre has been an enormous amounl of re- cent activily in biological weed con lrol in Au slra li a (Cullen a nd Delfosse 1990) for three main reasons: increasing f Hrmer -gra- lier and rural industry support; grea ter envi- ronmen tal awa reness by the population as a wh ole; and much-heightened interest by lo- ca l, St ate and Federal politi cians in non- chemica l pest control. Mos t of this ac tivity has centred around classica l biological con- trol, wh ere the targe t weed is from outside Aus trali a, and effective natural enemies of the weed are import ed from the weed 's na- live range . A major problem relati ng to weed man - agement in th e dried fruits industry is th at the weeds affecting the industry have not bee n determined. Th erefore, I propose three groups for weed s important to the industry: a small group of high-prior it y weed species in three ge nera (Emex, Tri bllius and Cenchms); a group of about 16 weed s peci es which are important targets for biological control for oth er r easons; and a group of about 50 weed species which are of occasional, regional or sporadic importan ce, some of which are al so future targets for biologica l control. To pl ace biologica l weed control in con- text , some of CS IRQis current projects are discussed briefly , including some weeds in all three categories. This is followed by recom- mendations on how such work cou ld proceed to maximize the benefit to the industry and to the science of biological control. A proposed breakdown of weeds of im- portance to the dried fruits industry One of Ih e most dimcull aspecls of deler- mining targe t weed species for bi ological con- trol for different commod it y or specia l inter- est groups is obtaining specific details on which weeds are import ant to each group. NlCr cons id erable checking, I could not find such a li sl for the dried fruils induslry. Therefor e, I propose to divide weeds of importance to the industry into three groups (Table I). This is obviously a slarling poinl only, and will change as new t echnology, chemica l and cultu ra l practices, etc., are in- trodu ced. Group One contains the three mos t im- portant genera of weed s to the dried fruits industry: 1.Emex spp., especi ally three-corn ered jack (E. olls/ralis Sleinheil, from South Africa; also kn own as spin y emex or do ubJegee) and lesscr Ja ck (E. spinosa [L.] ca mpdera , from northern Afri ca and western Eu- rope); 2. TribllllLfi /efTes/ris L. (puncturevi ne, from southern Africa and Mediterran ea n Eu- rope; also kn own as ca lt rop); and 3. Burr grasses (Cencllms spp., from NOrlh and Cenlral America), especia ll y ge nll e Annie (C. longispinll s [Hackel] Fernald). Group Two cont ains about 16 weed species Cfable I) whi ch are a lre ady lar ge ls for bio- logica l control because th ey are important environmentally, to ano th er commodity sec - tor, or are likely to be targets in the near fu- ture. Some of th ese, such as s kel eton weed (Chondrii/o jllncea L; Asteraceae) and Bathursl burr (Xonfhium spinosllm L. ; As- teraceae) ca n have a very severe impact on the industry; th e others are usuall y less im- porlanl. Group Three contains over 50 weed spe- cies which can occur heavily on a local, sca - sonal or sporadic basis. Many of th ese are Plant Proleclion Quarte rl y VO I.5 (3) 1990 91 potenlia ll argels for biol og i ca l control should funding become avai lable. Biological Weed Control Progra ms of Interest to the Dried Fruits Industry Group One Weed Species Three-cornered jack and caltrop. Spea kers at this Workshop have covered ecol ogy, chemical control and biological control of three-corner ed jack and caltrop in detail (Coo ke 1990, Bruzzese 1990, Gilbey 1990, Ma cGregor 1990, SCOII 1 99Oa,b, Shepherd 1990, Weiss 1990), and interested reade rs should refer to these p ape rs for further dc- tails on these topi cs. Spiny burr grasses, Cenchms s pp. (Pooceae ). There arc len Cencll",s spp. in Australi a, mostly from Central or North Amer ica, but there are al so two nat i ve spe- cies, and some benefi cia l pas ture speci es (Auld and Medd 1987). FUri her complicat- ing the situation is that the be neficial and native species ca n also be weedy. In addition to invading pastures, Cenchms spp. can be weedy because of the spines or burrs produced al flowering (Auld and Medd 1987). Considerable int erest was expressed by the Wool Resea rch and D eve lopment Fund (WRDF) for biol og i ca l conlrol of CenchnLS species, such as M ossman River grass, C. echinatus L. (an Ameri can bristly specie s), spiny burr gra ss or sa ndburr, C. in- cerWs M. Curtis, and spi ny burr grass or ge ntle Annie, C. longispinus (both Central American speCies). The WRDF funded a lil - erature review of the genus, which revealed a considerable amount of information on dis- tribution and control, but virtually nothing on potential biologica l control agent species. Dialogical control of CencJmls spp. would require foreign exploration in Ameri c.1 to find na tural enemies of the weeds and to study their ecology. This process would take a minimum of three years at a cos t of about $100,000 p.a. Group Two We ed Species Ske leton weed, Chondrilla juncea L. (Aster- aceae). The firs l use of a fun ga l phylO- pathogen , Puccinia chondrillina Bubak & Sydenham (U redinales), in bi ologi ca l co nl ro l was wilh Ihi s progra m (Cull en el aL 1973, Dclfosse el al. 1985). Three geneti ca!1y-dis- tin ct apomictic forms ofskclelOn we ed occur in Australi a. The introd uced strain of th e fun gus is eXlrcmely hosi-specifi c (onl y .1- taCking one of the forms of the weed), and stable (it is as virul en t now as it was when in - Iroduced nea rly 20 years ago). The program has a very hi gh benefil :cost ratio, giving tre- mendo us economic ret urns 10 the wh ea t in- dustry via the Wheat Research Council (WRC), which fu nd ed Ihe work (Cu!1en 1985, Marsden el al. 1980). The olher Iwo for ms of the weed are spreading into the ar- cas formerly dominated by the narrow-l eaf form, and current work centres arou nd fi nd-

Transcript of Biological control of weeds and the dried fruits industry 05-3 pp091-97 Delfosse.pdf · 14....

  • Biological control of weeds and the dried fruits industry

    Ernest S. Delfosse, CSIRO Division of Entomology, GPO Box 1700, Canberra, ACf 2601, Australia.

    Summary I propose that weed species of importance to the dried fruits industry can be divided into three groups. The first group contains the most important weeds to the industry: Emex spp. (especi~llly three-cornered jack, E. aus-tralis); puncturevine or caltrop (Tribulus ler-restTis); and gentle Annie lind related Cenchms spp. The second group contains weeds of lesser importnnce to the industry, hut because of their importance to other sec:tors of Australian agriculture and the environment, ure cu rrent targets for bio-logical control programs: skeleton weed (ChonLirilia jllncea) , Bathurst burr (Xant" -ium sp;nosllIn )j Noogoonl burr (X. slrumar;um)j thistles (CarduIlS, Cirsium, Onopordum and Silyhllln spp.)j 1)lIterson's curse (EcI,;um plan/ag;newn); com mon heliotrope (Helio/ropiutn europaewn)j St. John's wort (Hypericum perJoraJmn)j docks and sorrels (Rumex spp.) j and silverleaf nightshade (Solanum elaeagnifoliwn). The third group contains a large number of weeds of oct..'lsional, sporadic or geographi-cally-limited importance to the industry, some of which are turgets for biological con-trol programs in early stuges of develop-ment.

    Disclission of biological control pro-grams provides information on the types of weeds and biologicnl control Hgents cur-rently studied by CSIRO. Also discussed briefly are uspects of foreign exploration, opportunities for co-operation between Ru-ral Industry Research Funds, and co-opera-tion between CSIRO and the States in im-plementHtion of the programs.

    Introduction Thcre has been an enormous amounl of re-cent activily in biological weed conlrol in Auslralia (Cullen and Delfosse 1990) for three main reasons: increasing fHrmer -gra-lier and rural industry support; greater envi-ronmental awareness by the population as a whole; and much-heightened interest by lo-ca l, State and Federal politicians in non-chemical pest control. Most of this activity has centred around classica l biological con-trol, where the target weed is from outside Australia, and effective natural enemies of the weed are imported from the weed's na-live range.

    A major problem relating to weed man-agement in the dried fruits industry is that the weeds affecting the industry have not been determined. Therefore, I propose three groups for weeds important to the industry: a small group of high-priority weed species in

    three genera (Emex, Tribllius and Cenchms); a group of about 16 weed species which are important targets for biological control for other reasons; and a group of about 50 weed species which are of occasional, regional or sporadic importance, some of which are also future targets for biological control.

    To place biological weed control in con-text, some of CSIRQis current projects are discussed briefly, including some weeds in all three categories. This is followed by recom-mendations on how such work could proceed to maximize the benefit to the industry and to the science of biological control.

    A proposed breakdown of weeds of im-portance to the dried fruits industry One of Ihe most dimcull aspecls of deler-mining target weed species for biological con-trol for different commodity or special inter-est groups is obtaining specific details on which weeds are important to each group. NlCr considerable checking, I could not find such a lisl for the dried fruils induslry.

    Therefore, I propose to divide weeds of importance to the industry into three groups (Table I). This is obviously a slarling poinl only, and will change as new technology, chemica l and cultural practices, etc., are in-troduced.

    Group One contains the three most im-portant genera of weeds to the dried fruits industry: 1.Emex spp., especially three-cornered jack

    (E. olls/ralis Sleinheil, from South Africa; also known as spiny emex or doubJegee) and lesscr Jack (E. spinosa [L.] campdera, from northern Africa and western Eu-rope);

    2. TribllllLfi /efTes/ris L. (puncturevine, from southern Africa and Mediterranean Eu-rope; also known as ca lt rop); and

    3. Burr grasses (Cencllms spp., from NOrlh and Cenlral America), especia lly genlle Annie (C. longispinlls [Hackel] Fernald).

    Group Two contains about 16 weed species Cfable I) which are already largels for bio-logical control because they are important environmentally, to another commodity sec-tor, or are likely to be targets in the near fu-ture. Some of these, such as skeleton weed (Chondrii/o jllncea L; Asteraceae) and Bathursl burr (Xonfhium spinosllm L.; As-teraceae) can have a very severe impact on the industry; the others are usually less im-porlanl.

    Group Three contains over 50 weed spe-cies which can occur heavily on a local, sca-sonal or sporadic basis. Many of these are

    Plant Proleclion Quarterly VOI.5(3) 1990 91

    potenliallargels for biological control should funding become avai lable.

    Biological Weed Control Programs of Interest to the Dried Fruits Industry

    Group One Weed Species Three-cornered jack and caltrop. Speakers at this Workshop have covered ecology, chemical control and biological cont rol of three-cornered jack and caltrop in detail (Cooke 1990, Bruzzese 1990, Gilbey 1990, MacGregor 1990, SCOII 199Oa,b, Shepherd 1990, Weiss 1990), and interested readers should refer to these papers for further dc-tails on these topics.

    Spiny burr grasses, Cenchms spp. (Pooceae). There arc len Cencll",s spp. in Austra lia, mostly from Central or North America, but there are also two nat ive spe-cies, and some beneficia l pasture species (Auld and Medd 1987). FUriher complicat-ing the situation is that the beneficial and native species can also be weedy.

    In addition to invading pastures, Cenchms spp. can be weedy because of the spines or burrs produced al flowering (Auld and Medd 1987). Considerable interest was expressed by the Wool Research and Development Fund (WRDF) for biological conlrol of CenchnLS species, such as M ossman River grass, C. echinatus L. (an American bristly species), spiny burr grass or sandburr, C. in-cerWs M. Curtis, and spiny burr grass or gentle Annie, C. longispinus (both Central American speCies). The WRDF funded a lil -erature review of the genus, which revealed a considerable amount of information on dis-tribution and control, but virtually nothing on potential biological control agent species.

    Dialogical control of CencJmls spp. would require foreign exploration in Americ.1 to find natural enemies of the weeds and to study their ecology. This process would take a minimum of three years at a cost of about $100,000 p.a.

    Group Two Weed Species Skeleton weed, Chondrilla juncea L. (Aster-aceae). The firs l use of a fungal phylO-pathogen, Puccinia chondrillina Bubak & Sydenham (Uredinales), in biological conl rol was wilh Ihis program (Cullen el aL 1973, Dclfosse el al. 1985). Three genetica!1y-dis-tinct apomictic forms ofskclelOn weed occur in Australia. The introduced strain of the fungus is eXlrcmely hosi-specific (only .1-taCking one of the forms of the weed), and stable (it is as virulent now as it was when in-Iroduced nearly 20 years ago). The program has a very high benefil :cost ratio, giving tre-mendous economic returns 10 the wheat in-dustry via the Wheat Research Council (WRC), which fu nded Ihe work (Cu!1en 1985, Marsden el al. 1980). The olher Iwo forms of the weed are spreading into the ar-cas formerly dominated by the narrow-leaf form, and current work centres around find-

  • 92 Plant Protection Quarterly VOI.5(3) 1990

    ing strains of the fungus in Europe which arc virulent against these two other forms of the weed.

    Othcr agent specics havc bcen released ag,ainst skeleton weed (Julien 1987), but only the rust fungus contributes significant ly to its control. Current work in Europe has charac-terized over 300 forms o f the weed, and has dcfincd two arcas in wcstcrn Turkey which have the highcst levc l of variability and offer the best ehflnce of obtaining additional useful strains of P. cJlOnd,.i//ina. Scveral new strains of the fungus which are virulent against the remaining two forms of t he weed have been found, but none arc as virulent as the initial strain for the narrow-leaf form (J.M. Cullen, personal communication, 1990). U~lthurst burr, Xanlllium spinosllln L. (As-teraceae). This South American species was included, inappropriately, in th~ initial inves-tigations on Noogoora burr in North Amer-ica, where it is an introduced weed. No in-sects have been introduced deliberately for classical biological control of this species. However, an accidentally-introduced seed ny, Ellaresla bllllam' Wicdemann (Diptera: Tephritidae) was first noted in 1918. It is widely established and destroys large num-bers of seeds, but seems to have no effect on the overall distribution and abundance of the weed (Wilson 1960).

    Recently, Bathurst burr has been the tar-get of an augmentative/inundative biological control program whcre the aim is to develop a mycoherbicide from an indigenous pathogen, Collelotrichwn x{lnlhii Halsted (Melanconiaceae), known to be capable of extensively damaging the weed in the field. It was first reported by Butler (1951). Com-mercial production of this promising pathO-gen is current ly being investigated (Auld ef al. 1986). Natural enemies for Bathurst burr will also be investigmed soon HS pan of a new CSIRO/New South Wales Department of Agriculture and Fisheries initiative, funded by the WRDF. Noogoora burr, Xanlltiwn sl17lmariwn L. (Asleraceae). This intractable species was one of the first target species for biological control in Australia, staning in 1929 (Wilson 1%0, julien 1987, Wapshere 1974). Several agent species were released, including: a seed fiy species, Euaresta aeqllalis Loew (Diptera : Trypetidae) from North America in 1932 (which established, but provides no control); two beetle species, Mecas satllrnina Le Conte (Coleoptera: Cerambycidae) from North America in 1963, and Nupserlw vex-ator (Pascoe) (Coleoptera: Cerambycidae) from India in 1964 (both species established but neither provides any significant control); an accidentally-introduced rust fungus, PflC-cinia ;Wnl!iii Schweinitz (Uredinales), found in 1974, before proJXlsed work on it as an agent species was carried out (very damaging to the weed under particular conditions, but effectiveness varies from season-to-season and from region-to-region; Julien el al.

    Table 1. Priority weeds to the dried fruits industry.

    Weed species Country of origin

    Group 1 (p riority weeds to Ihe dried fruits induslry for biological control) I. Three-cornered jacks (Emex australis Steinheil; South Africa

    POlygonaccac) 2. Lesser jack (E. spinosa [L.] Campdera; POlygonaceae)

    3. Caltrop (Tribulus ten-estris L.; Zygophyllaceae)

    northern Africa and western Europe southern Africa and Mediterra-nean Europe

    Spiny burr grasses (Asteraceae; ten weedy Cenchrus spp. in Australia), especially 4. Mossman River grass (C ecllinaf1IS L.) North America 5. Spiny burr grass or sandburr (C incerlus M. Curtis) Central America 6. Gentle Annie (Clongispimls [Hackel] Fernald) Central America

    Group 2 (important weeds to the industry which are or have been targets for biological control) 1. Skeleton weed (Chondrilla jllncea L.; A~teraeeae) 2. Bathurst burr (Xantltiwn spinoswn L; Asteraceae) 3. Noogoora burr (X stnllnariwn L) 4. Scotch and related thist les (Onopordwn. spp.;

    A'Steraceae)

    Transcaspian Central Europe Chilc North America Mediterranean Europe

    5. Slender and nodding thistles (Cardllus spp.; Asteraceae) Mediterranean Europe 6. Spear thistle (Cirsiwn vlligare [Savi] Tenore; Mediterranean Europe

    Asteraceae) 7. Variegated thistle (Silybwn marianwn. [LI

    Gaertncr; Asteraceae) Mediterranean Europe

    8. Slender and nodding thistles (CarcifUls spp.; Asteraceae) Mediterranean Europe 9. Paterson's curse/salvation Jane (Echiwn Mediterranean E urope

    piantagineuln L.; Boraginaceae) 10. Common heliotrope (Heliolropiflm europaewn L;

    8oraginaeeae) II. S1. John's wort (Hypericum perfomtwn L.; Clusiaceae) 12. Horehound (Man'lIbiwn vulgare L; Lamiaceae) 13. Docks and sorre ls (Rumex spp.; POlygonaccae) 14. Silverleaf nightshade (Solanum elaeagnifoliwn

    Cavanilles; Solanaceae)

    Mediterranean Europe to North Nrica Mediterranean Europe Mediterranean Europe Europe Central and South America

    Group 3 (important weeds to the industry which a re not currently targets for biological control) 1. African lovegrass (Eragrostis CIl11'II/a ISchrader]

    Nees; Poaceae) 2. Amaranth (Amaranllills spp.; Amaranthaceae)

    3. Barley grasses (Hordeum spp.; Poaceae) 4. Barnyard grass (Eehinochloa ems-galli [L];

    Beauvisage; Poaceae)

    South Africa

    tropical and subtropical America Europe and Asia Europe and India

    5. Blackberry nightshade (So/anum nigmm L ; Solanaceae) CosmoJXllitan 6. Burr marigold (Ridens subalternana DC.; Asteraceae) America 7. Capeweed (Aretot"eea calendula [L.] South Nrica

    Levyns.; Asteraceae) 8. Cobbler's pegs (Eidens sllballemana DC.; Asteraceae) America 9. Common evening primrose (Oenor/tera slricta South America

    Ledcbour ex Link; Onagraceae) 10. Couch grass (Cynodon dactylon [L.]; Persoon; Poaceae) Cosmopolitan 11. Fat hen (Chenopodium album L; ChenoJXldiaceae) European and Asian (possibly

    12. Field bindweed (ConvotVlI/US arvensis L; Convolvulaceae)

    13. Fumitory (I'II/naria spp.; Fumariaceae) 14. Johnson grass (Sorghum l/alepense [L.] Persoon;

    Poaceae) 15. Kikuyu (Pennisetwn clandestinwn. Hochstetter

    ex Chiovenda; Poaceae) 16. Mallow (Malva 'pp.; Malvaceae) 17. Mcdics (Medicago spp.; Fabaceac) 18. Morning glory (Ipomoea spp.; Convolvulaceae) 19. Nutgrass (Cypems rotlmdlis L.; Poaceae)

    cosmopolitan) Europe

    Europe Mediterranean region

    Africa

    Mediterranean Mediterranean tropical Cosmopolitan

    continued

  • Table 1. I'riority weeds to the dried fruits industry (continued).

    Weed species

    20. Onion weed (A.\p/iodelus jistlll.oslts L.; Liliaceae) 21. Oxahs and sorrels (Oxalis spp.; Oxahdaceae)

    Country of origin

    Mediterranean Europe South Africa, South America and cosmopolitan spp.

    22. Paspalum (Paspnlum dilatatIOn Poiret; Poaceac) South America 23. Perennial thistle (Cirsillln. an'cnse IL.] Scopoli; Europe

    Asteraceae) 24. Phalaris (Pllalaris aqua/ica L.; Poaceae) 25. Pigcon grass (Setaria spp.; Pooceae) 26. Prickly lettuce (Lnctllca se'Tiola L.; Asteraceae)

    Mediterranean region Central and South America Europe

    27. Rhodes grass (C"'oris gayana Kunth; Poaceae) tropical America 28. Silverleaf nightshade (Solanum e/aeagnifoliwn Central and South America

    Cavanillcs; Solanaccae) 29. Soursob (OXI/lis pes-caprae L.; Oxahdaceae) 30. Stinging nettle (Urtica spp.; U rticaceae)

    Sout h Africa Europe

    31. Stinkweed (NGI·arrelia squan·osa [Eschscholtz] H ooker & Arnott ; Polemoniaceae)

    North America

    32. Subterranean clover (Trifolium subte"onewn L.; Fabaceae)

    Europe

    33. Thistles, annual (Card flll s, Cirsium, Onopordwn, Europe Silybwn spp.; Asteraceae)

    34. Thornapples (Datura spp.; Solanaceae) 35. White clover (Trifolium repens L.; Fabaceae)

    America Europe Australia 36. Willow herb (Epilobimn billnrdierinnwn Seringe;

    Onagraceae) 37. Wild oats (Awna fallta L.; Poaceae) 38. Wild radish (Rnplulnus raplwnistlion L.; Brassicaceae)

    Medi terranean region Europe Mediterranean region Cosmopolitan

    39. Wimmera ryegrass (Lolium rigidwn Gaudin; Poaceae) 40. Wireweed (Polygonmn aviclllnre L.; POlygonaceae) 41. Yellow burr weeds (Amsinckia spp.; 8oraginaceae) North and Central America

    Unspecified species 42. Annual broad leaf weeds 43. Annual grasses 44. Perennial broad leaf weeds 45. Perennial grasses

    1979); and a moth, Epiblema slrenllana (Walker) (Lepidoptera: Tortricidae) in 1984 (introduced for control of annual ragweed and parthenium weed, but which also pro-duces considerable damage to Noogoora burr). Thus over the last to to 15 years, dam-age to the weed (particu larly by P. xnnthii in tropical and subtropical areas) has increased significantly and its importance has declined in some areas, but further biological control agent species are certainly required to pro-duce beller control. Saffron thistle, Cart!tamus lnnaws L. (A~teraceae) Detailed allcn tion has not yet been given to saffron thistle, but work on this spe-cies is planned in the long term. Slender ~tnd nodding thistles, CardllliS spp. (Asteraceae). Nodding thistle, CardtlliS manns L., and slender thistles, C. pycno-cep!tallls Land C. tenlliflo17ls Curtis, were the first thistle species examined in detail , commencing in 1986.

    Nodding thistle has been successfully con-trolled with natural enemies in other parts of the world, so Australian studies concentrated initially on the role of natural enemies in regulating the weed's popUlation in Europe and on the biology of the most significant species (Sheppard el al. 1989). So far two

    many countries many countries many countrics many countries

    agent species have been introduced to Aus-tralia: three populations of a seed-head weevil , Rhinocyllus conicus Froehlich (Cole-optera: Curculionidae), released in 1988-89 at sites on the southern and northern Table-lands of New South Wales; and a seed ny, Urophora solrli/ialis (L.) (Diptera: Tephriti-dae), which was imported into quarantine in 1990 for host-specificity testing. Studies are in progress on other natural enemies of nod-ding thistle.

    A population of R. coniclls which attacks the slender thistles and variegated thistle, Si-lybwn marianun (L) Gaertner, has been in-troduced and release in Victoria, in coopera-tion with the Victorian Department of Con-serva tion and Environment (VDC&E). A weevil, Cemorhync/ws trimaculalllS (F.) (Celeoptera: Curculionidae), is being evalu-ated in Europe, and a search is being made for virulent strains o f a rust fungus which is already present in Australia, Puccinia cardui-pycnoceplwli H. SydOW & I'. SydOW (Uredi-nales). Variegated thistle. So far, the only agent species to be released for variegated thistle is the type of R. conicliS which also auacks slen-der thistle, as mentioned above. Detailed at-tention has nOl yet been given to this species,

    Plant Protection Quarterly VOl.5(3) 1990 93

    but work is planned in the long term. Spear thistle, Cirsiwn vulgare (Savi) Tenore. Surveys for potential biological control agents are underway on this species in Eu-rope. along with surveys on other thistle spe-cies. A population of R. coniclls for spear thistle has been imported into quarantine from southwest France, and application has been made for importation ofa seed fly, Uro-phoro slylalo (F.) (Diptera: Tephritidae). Scotch and related thistles, Onopordwn spp. (Astraceae). Detailed st udy of Scotch thistle, Onopordwn ncanlhiwn. L., lIIyrian thistle, 0. il/yricwn L., and stemless thistle, 0. acaulon L ,started in 1987-88. Tn progress are population studies on TIIyrian thistle in Australia, and assessmcnt of several natural enemies in E urope. These include: seed-a t-taCking weevils, Larin/iS cynareae F. and L lahlS (Coleoptcra: Curculionidae) (the latter species is now in quarantine in Australia for host-speCificity testing), seed-attacking nics, Tephritis posticn Locw and Terellia gyneco-chroma H ering (Diptera: Tephrilidae); a stem-boring beetle, Loots cnrdui Olivier (Coleoptera: Curculionidae); and a sap-feed-ing bug, Tettigometra supllllra Mulsant (Hemiptera: Tettigometridae). Paterson's curse and related Echimn spp. (Boraginaceae). 8iological control of £Chillm spp. was delayed from 10 July 1980 to 17 November 1988 by a High Ceurt in-junction granted on behalf of two beekeepers and two graziers. Because of this conOict-of-interest. it is probably the best known current CSIRO project. This delay cost the Austra-lian community more than $300 million in direct losses, and much more in indirect losses. SpeCific legal aspects of this program have been documented (Cullen and Delfosse 1985, Delfosse 1985a, Delfosse and Cullen 1981b) and will not be discussed here.

    The first agent species to be imported against Paterson's curse since the legal con-flict was resolved is a leaf-mining moth, Din· leclica scalariella (Zcllcr) (Lepidoptera: Gracillariidae) (Cullen and Delfosse 1990). This species was released in mid-1980, but did not establish due to a combination of low numbers and premature removal of the host plant that season due to drought and grass-hopper plagues (Delfosse el al. 1987). As a result of large releases by the States in this cooperative program, D. scalariel/a is widely established, spreading rapidly, and extremely large numbers have been recorded in some areas. The critical point will be the popula-tion level it eventually attains and is able to maintain, particularly if attacked by native parasites.

    There are at least twenty agent species available for Echiwn. Some of the higher-rated approved species are: two rosette- and root-attacking weevils, Ceworhynclms larva-ilL< Schultze and C. geogmphiclls (Goeze) (Coleoptcra: Curculionidae); twO rosette-and root-allacking nea beetles, LongiwrsllS ecllii Koch and L. aeneus KUlsch (Coleop-

  • 94 Plant Protection QUClrterly V OI. S(3) (990

    tera: Chrysomelidae); two cell-sucking bugs, Diclyla nassata PUlon and D. eellii Schrank (Hemiptera: Tingidae); and Cl stem-boring beetle, Phytoecia coem/escens (Scopoli) (Coleoptera: Cerambycidac).

    Small numbers of C. lOll/an IS were re-leased at one site in mid- 1989. A current de-lay is the lack of an efficient rearing system for Ihis difficult species, but further releases are expected in 1990-9 1. L. aenells is also being reared under quarantine conditions. Other species to be tested include: Cl rosette-and bud-allacking mot h, EI/llnia bipllnctella F. (Lepidoptera: Ethmiidae); Elhmia ter-minella Fletcher, whose larvae attack stem buds; two species of nower beet les, Meli-gellies planillscu/liS HeeL and M. Iristis Sturn. (Coleoptera: Nitidulidae); an eriophyid mite, probably Eriopliyes eellii Canestrini (Acari: Eriophyidae), and several pathogens.

    With the wide interest in this program, there is extensive collaboration between CSIRO and all State D epartments, and an intensive cooperat ive program of release, moni toring and evaluation is underway. Eu-ropean studies on the ecclogy of the plal1l and its natural enemies arc on-going. Common heliotrope, Heliotropiwn ell-ropaell1n L. (Doraginaceae). This summer-growing, annual M editerranean weed (Delfosse and Cu llen 1981a) ca uses damage in excess of $46 million p.a. (Delfosse and Cullen, unpubl. data). The unreliable, Hlmost ephemeral occurrence of this species from season-Io-season and from paddock-to-pad-dock suggests that it would be a difficult tar-get for classica l biological control with ar-th ropods.

    A flea beetle species, L ongiwrsl/s albineus (Foudras) (Coleoptera: Chrysomelidae), was released several times from 1979 to 1989. Though recoveries have been made at east-ern sites, results have been disappointing. Further European and Australian ecological studies of soil type, fertility and plant growth are underway to try to explain the insect's unexpected poor performance (Delfosse 1985b).

    Two more common heliotrope agents being prepared for release arc a leaf-attack-ing rust fungu s, Uromyees helio fropii Sredinski (Uredinales) and a fOliage- and root-allacking weevil , Pachycerus cordiger Gcrmar (Coleoptera: Curculionidae). The rust is extremely damaging to common helio-trope in Europe, causing death of the plant and massive reduction of seeding (Hasan 1985). A proposal for release has been made. Adults of P. cordiger feed on common helio-trope fOliage, and larvae feed on roots. IL at-tacks a wide range of Doraginaceae under cage conditions (Huber and V ayssieres 1990), but is more restricted in the field; test-ing of Australian native n oraginaceae under field conditions in Europe, the first lime this was done in an Australian program, resulted in a decision to apply for its release for sum-mer 1990-91.

    Two other potential agent species are being studied: a bud-feeding moth, £I/llnia disligmarella Erschoff (Lepidoptera: E th-miidae), for which host-specifici ty tests will begin in quarantine in Australia as soon as a colony can be established: and another fun-gal pathogen, Cercospora he/iotropii-boceoni Scalia (I-Iyphomycetcs), for which tests have begun at the CSIRO Biologica l Control Unit in Montpellier, France.

    Since 1979 there has been extensive col-laborat ion between CSIRO and relevant State Departments on this program. In par-ticu lar, in 1988 a cooperative program of re-lease, monitoring and evaluation of agents with CSIRO and the Western Australian Department of Agriculture (WADA) was started. St. John's wort, Hypericum pelfomlwn. L. (Clusiaceae). The current program aims to improve the level of control exerted by the beet le Clllysolina qllatirigemina (Suffrian) (Coleoptera: Chrysomelidae) imported in thc 1940s (Delfosse and Cullen 1981c). Re-cent releases include: new populations of C. qlladrigemina; a rOliage-feeding moth species, Anaitis efformata Guence (Lepidop-tera: Geometridae), whieh failed to establish largely due to ant predation of larvac; an-other moth, A ctinotia hyperici Schiff. (Lepi-doptera: Noctuidae), which has not been re-covered in the field; an aphid, Aphis chloris Koch (Hemiptera: Aphididae), which is well-established and spreading; and a new popula-tion of Agrillls hyperici (Creuzer) (Coleop-tera: Duprestidae), wb.ich is being evaluated (Briese 1986, 1989). The most promising agent species is currently in quarantine in Canberra. It is a mite, A culus hyperici (Liro)(Acari :Eriophyidac), which we hope to release this year. Horehound, Mamlbiwn vllfgare L. (Lamiaceae). A preliminary survey for natu-ral enemies of horehound was conducted in Europe by CSrRO several years ago, and several potential agents were identified. Cur-rently, there is renewed interest in this weed due ( 0 problems it is causing in conservation areas, and a detailed program will commence later this year as a cooperative program be-tween CS1RO and the VDC&E. Docks ~md sorrel, Rwnex spp. (Polygo-naceae). Several dock species (Rllmex spp.) plus sorrel (R. acelosella L) are weeds in several States, particularly R. pulcher L. in Western Australia. A W ADA-CSlRO proj-ect has identified two highly-promising moth species in Europe: Bembecia c/llysidiformis (Esper) and Chamae~phaecia dory"formis (Ochsenhcimer) (Lepidoptera: Sesiidae), whose larvae bore into the perennial root-stock of mature plants, often killing them. Both species have been imported and large numbers of C. dorylijormis were released in summer 1989-90, with initial indications of high surviva l rates. Further work will concen-trate on the second species, followed, if nec-essary, by two other potential agent species,

    Perapion vio/acelIIn Kirby (Coleoptera: Api-onidae) and Pegomyia nigritarSlls Zett. (Diptera: Anthomyidae). Silverleaf nightshade, Solanum elaeagnifo-lilUn. Cavanilles (Solanaceae). Biological con-trol of silverleaf nightshade has been re-quired for a number of years. Studies have been made in Australia, the USA and South Africa (Field et al. t988, Orr et al. 1975, Robinson et al. 1978, Zimmermann 1974). Apart from problems of lack of specificity of several possible agents (Siebert 1975, 1977), there was little prospect of success, mainly because most natural enemies of silverleaf nightshade are adapted to the presumed area of origin of the plant in north-eastern Mex-ico, and are climatically unsuited to the re-gions of Australia infested by the weed (Wapshere 1988).

    The potential agent species with the high-est priority is a nematode, Om·nia phyllobia (Thorne) (Nematoda: Neotylenchidae), which has been the SUbject of detailed study in a joint project of the Victorian Depart-ments of Agriculture and Rural Affairs, and Conservation and Environment. The nema-tode forms galls on silverleaf njghtsbade, which weaken and can gradually kill the plant. Studies of its host-specificity have demonstrated that under certain conditions, galls can be formed on egg plant, Solanum me/ongena L., and on some native species of So/anum. The risk that this poses is currently being evaluated (R. Field, pers. comm., 1990).

    GrOllI' Three Weed Species: Fumre Bio-logical Control Programs of POIential Interest 10 Ihe Dried Fnlils Industty M ost problem weeds in Australia have been proposed as targets for biological control, but only a proportion have so far been tackled. A few important species have not yet been con-sidered in any detail or are only at a very pre-liminary stage of investigation. For example, there are over 40 current biological control of weeds projects in Australia (Cullen and Delfosse 1990), most of which involve only classical/inoculative biological control, where both the pest weed and its natural enemies are exotic (Wapshere et at. 1989), and there arc many more in earlier stages of develop-ment.

    It is important to note three things: l ) ln contrast to earlier times, many of these

    projects are collaborative ventures be-tween CSIRO and State groups;

    2)Australia has only onc current augmenta-tive/inundative mycoherbicide project, no programs involving conservation of natu-ral enemies, and no programs involving grass weeds as targets for biological con· trol; and

    3)AlI of the following programs are funded to a very large degree by Rural Industry Research Funds (RIRFs), particularly the WRC, the Australian Meat and Livestock

  • Research and Development Corporation (AMLRDC) and the WRDF. Without these funds, the research could nOt be con-ducted.

    Some potential targets for biological control include: Onion weed, Asphodellls fisndosus L. (LiIi-aceae). Onion weed has never been the sub-ject of detailed surveys, but a damaging rust fungus, Pllccinia barbeyi (Roum.) Magn. (Uredinales) was found to have some poten-tial during preliminary surveys of this plant in Europe. The biology has now been eluci-dated and this rusl seems to be host-specific and quite damaging. Capeweed, Arctothecn calendllla (L.) Lcvyns. (Astcraceae). This specics has often been proposed as a possible target for bio-logica l con trol, but only in eastern Australia, where it causes considerable losses (Sloan et aL 1989). In Western Australia however, al-though recognized as a problem, it has a more ambiva lent status, many farmers con-sider it as va luable feed for stOCk, and call it "capcfeed" ! While tbis precludes a biOlogi-cal control program in the short term, a pos-sible change in attitude in the long term would allow a program to proceed. A pre-liminary survey of capeweed in South Africa has revealed some promising natural ene-mies (SCOlt and Way 1990), but further work cannOl be justified at present. Soursob, Oxlliis pes-cnprae L (Oxalidaceae). Very preliminary surveys of this weed in South Africa several years ago seemed to indicate a lack of potential agents, but more recent work has shown the existence of sev-eral promising species. A more detailed proj-ect is expected to commence in the near fu-ture. Yellow burr weeds, Amsinckia spp. (Barag-inaceae). There have been varying assess-ments of the importancc of these American weeds, with some States rming them highcr than others. While low in priority at prescnt, surveys have been made of these weeds in O:l lifornia, and potential agents have been identified (Detfosse, unpubl. data). Gruss weeds. There has been no work on classical biological control of grass weeds in Australia, but the potential for their control has recently been reviewed (Wapshere 1990). There is some potential for classical biological control of certain species, which should be investigated in more detail. These include grasses of the genera Bromm, H O/CIIS , Ecl!inoclllon , Nasel/a , Cortaderin, Elellsine and Rotfboellia , piUS Johnsongrass, Sorg""'" Iwlepense (L.) I'ersoon (Massein and Lindow 1986, McFayden 1985).

    Despite the above reference to classical biological control of grass weeds, currently, the only acceptable approach to these species would be by using mycoherbicides. Work is currently in progress to investigate the poten-tial of mycoherbicides for control of several annual grass weeds by infection of the seeds while in the soil (Medd eI al. 1986). The im-

    partance of several grass weeds in pastures, their ambiva lcnt stat us in certain areas and thc question of specifiCity of agent species where the weed is very closely-related to im-portant pasture, crop and native species, would all suggest that the pOlential for myeo-herbicides should be investigated in consid-erably more detail. While repeated applica-tion of a mycohcrbicide, with its associated cost , is necessary, the very fact that its effect is limited in time and space allows the avoid-ance of connict-o f-intcrest problems, whether concerning the target weed in differ-ent situations or related non-target species. Potential augmentative/inundalive pro-grams. This area has grem potential for de-velopment. Current research is qui te limited, the only group inVOlved concentrating on Bathurst burr and the seeds of grass weeds. In this commercial field , the size of markets will playa major role in determining targets (Wapshere 1987). Even though the Austra-lian market will be relatively small, increasing restrictions on herbicide use, pressure from environmental lobby groups, and increase in wced resistance to herbicides could all force further development. N~ltive woody weeds. Cullen and Delfosse ( 1990) concluded that native woody weeds, which have become a major problem in the pastoral zone of inland New South Wales and Queensland, are potential targets for augmentative/inundative biological control, but not generally for classica l biological con-troL They all already have cocvolved natural enemies in Australia, but these arc ineffec-tive because of environmental f

  • 96 Plant Protection Quarterly VOI.5(3) 1990

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