Citation: Zachariades, C. (2018) Biological control of invasive alien plants in South Africa: a list of all , mites and pathogens released as biological control agents from 1913- 2018. http://www.arc.agric.za/arc-ppri/Documents/Table2-NaturalEnemiesReleased.pdf

Table 2. Target weed species in South Africa, their region of origin and degree of biological control achieved (see below for definitions): natural enemies released, or for which a release permit has been obtained, their feeding guilds or association with the host plant, year of release, establishment, and degree of agent damage to the weed (if established) (see below for definitions), and key references for each (see separate references document). Winston et al. (2014) was consulted for some sources of agents.

Scientific name and author Degree of Natural enemy Feeding guild Agent research Damage inflicted Source of agent Key references Common name control status Region of origin ARACEAE Pistia stratiotes L. Complete Neohydronomus affinis Hustache Leaf and stem Released 1985, Extensive Brazil via Australia Cilliers 1991; Coetzee et water lettuce (Coleoptera: Curculionidae) borer established al 2011b South America ASTERACEAE Ageratina adenophora (Spreng.) Negligible Passalora ageratinae Crous & A.R. Leaf spot Released 1987, Moderate Prob. Central Morris 1989, 1991; R.M.King & H.Rob. (= Eupatorium Wood pathogen established America, prob. via Buccellato et al. 2012 adenophorum Spreng.) (=“Phaeoramularia” sp.) Hawaii (USA), via crofton weed (anamorphic fungus; Australia Mycosphaerellales: Central America Mycosphaerellaceae)

Procecidochares utilis Stone (Diptera: Stem galler Released 1984, Moderate Mexico Kluge 1991; Erasmus et Tephritidae) established al.1992; Buccellato et al. 2012 Ageratina riparia (Regel) R.M.King & Complete Entyloma ageratinae R.W. Barreto Leaf pathogen Released 1989, Considerable Jamaica via Hawaii Morris 1991 H.Rob. (= Eupatorium riparium &H.C. Evans (Entylomatales: established (USA) Regel) Entylomataceae) mistflower Central America Campuloclinium macrocephalum Negligible Cochylis campuloclinium Brown Flower feeder Release permit Argentina Brown 2006; (Less.) DC. (formerly Eupatorium (: Tortricidae) issued McConnachie et al. 2011 macrocephalum Less.) pompom weed Liothrips tractabilis Mound & Pereyra Stem galler Released 2013, Moderate Argentina Mound & Pereyra 2008; South America (Thysanoptera: Thripidae) established McConnachie et al. 2011

Puccinia eupatorii Dietel (Pucciniales: Leaf rust Not released, occurs Moderate McConnachie et al. 2011; Pucciniaceae) pathogen locally Retief et al. 2016

Chromolaena odorata (L.) R.M.King Substantial Calycomyza eupatorivora Spencer Leaf miner Released 2003, Trivial Jamaica Zachariades et al. 2002; & H. Rob. (formerly Eupatorium (Diptera: Agromyzidae) established Strathie et al. 2007; odoratum L.) Zachariades et al. 2011b; triffid weed Nzama et al. 2014a&b Dichrorampha odorata Brown & Stem-tip galler Released 2013, Jamaica Brown & Zachariades North, Central & South America Zachariades (Lepidoptera: Tortricidae) establishment 2007, Dube et al. 2017 unconfirmed Lixus aemulus Petri (Coleoptera: Stem borer Released 2011, Acre state, Brazil Zachariades et al. 2002; Curculionidae) establishment Kluge & Zachariades 2006 unconfirmed aurata (Butler) Leaf feeder Released 1990, not North-western Kluge & Caldwell 1993a; (Lepidoptera: ) established Argentina Zachariades et al. 1999

Pareuchaetes insulata (Walker) Leaf feeder Released 2001, Extensive, Florida, USA; Kluge & Caldwell 1993b; (Lepidoptera: Erebidae) established localised Jamaica; Cuba Strathie & Zachariades 2004; Strathie et al. 2007; Zachariades et al. 2011b; Dube et al 2009, 2014; Uyi et al. 2014 a&b, 2015, 2016a&b, 2017, Zachariades et al. 2016 Pareuchaetes pseudoinsulata Rego Leaf feeder Released 1989 & Trinidad via USA; Strathie & Zachariades Barros (Lepidoptera: Erebidae) 1998, not Trinidad via 2002; Zachariades et al. established Indonesia; Northern 2011b Venezuela Recchia parvula (Lane) Stem and crown Released 2016, North-western Zachariades et al. 2007; (Coleoptera: Cerambycidae) borer establishment Argentina Zachariades et al. 2011b unconfirmed Cirsium vulgare (Savi) Ten. Negligible Rhinocyllus conicus (Froelich) Seed feeder Released Moderate France Zimmermann 1991; spear thistle (Coleoptera: Curculionidae) 1984,1986, Hodson et al. 2003 established Europe Urophora stylata (Fabricius) (Diptera: Seed feeder Released 1984, Germany, France Zimmermann 1991 Tephritidae) 1987, 1989, 1990, not established Parthenium hysterophorus L. Under Listronotus setosipennis (Hustache) Stem borer Released 2013, Considerable Argentina Dhileepan & Strathie parthenium evaluation (Coleoptera: Curculionidae) established 2009; Strathie et al. 2011

Caribbean Puccinia abrupta var. partheniicola Leaf rust Not released, occurs Moderate Unknown origin Wood & Scholler 2002; (Jack. & Holw.) (Pucciniales: pathogen locally Dhileepan & Strathie 2009 Pucciniaceae)

Puccinia xanthii Schwein. var. Leaf rust Released 2010, Considerable Australia (originally Dhileepan & Strathie parthenii-hysterophorae Seier, H.C. pathogen established Mexico) 2009; Strathie et al. 2011 Evans & Á. Romero (=P.melampodii; Pucciniales: Pucciniaceae)

Smicronyx lutulentus Dietz Seed feeder Released 2015, Unknown Australia (originally Dhileepan & Strathie (Coleoptera: Curculionidae) established. Mexico) 2009; Strathie et al. 2011

Zygogramma bicolorata Pallister Leaf feeder Released 2013, Considerable Australia (originally Dhileepan & Strathie (Coleoptera: Chrysomelidae: established Mexico) 2009; Strathie et al. 2011; Cassidinae) McConnachie 2015a; Chidawanyika et al. 2017 Silybum marianum (L.) Gaertn. Not Rhinocyllus conicus (Froelich) Seed feeder Alternative host, California, USA Zimmermann 1991 milk thistle determined (Coleoptera: Curculionidae) released 1985, not established Mediterranean, Asia Tithonia diversifolia (Hemsl.) A.Gray Not Physonota maculiventris (Coleoptera: Leaf feeder Release permit Mexico Mphephu et al. 2017 Mexican sunflower applicable Chrysomelidae: Cassidinae) issued

Central America Tithonia rotundifolia (Mill.) S.F.Blake Under Zygogramma piceicollis (Stål) Leaf feeder Released 2015, Considerable, Mexico Simelane et al. 2011b red sunflower evaluation (Coleoptera: Chrysomelidae: established localised Chrysomelinae) Central America Zygogramma signatipennis (Stål) Leaf feeder Released 2015, Considerable, Mexico Simelane et al. 2011b (Coleoptera: Chrysomelidae: established localised Chrysomelinae)

AZOLLACEAE Azolla cristata Kaulf. Complete Stenopelmus rufinasus Gyllenhal Frond feeder Alternative host, Extensive USA Madeira et al. 2016 Carolina water fern (Coleoptera: Curculionidae) released 1997, established North and South America Azolla filiculoides Lam. Complete Stenopelmus rufinasus Gyllenhal Frond feeder Released 1997, Extensive USA McConnachie et al.2004; red water fern (Coleoptera: Curculionidae) established Hill et al. 2008; Coetzee et al. 2011b South America BASELLACEAE Anredera cordifolia (Ten.) Steenis Under Plectonycha correntina Lacordaire Leaf feeder Released 2016, Trivial Buenos Aires, Cagnotti et al. 2007; Van subsp. cordifolia evaluation (Coleoptera: Chrysomelidae: established Argentina der Westhuizen 2011 Madeira vine Chrysomelinae)

South America BIGNONIACEAE Dolichandra unguis-cati (L.) Lohman Negligible Carvalhotingis hollandi Drake Leaf sucker Released 2007, Unknown Misiones, Argentina Williams et al. 2008 (formerly Macfadyena unguis-cati (Hemiptera: Tingidae) established (L.) A.H.Gentry) cat's claw creeper Carvalhotingis visenda Drake & Leaf sucker Released 2007, Moderate Misiones, Argentina Williams et al. 2008 Hambleton (Hemiptera: Tingidae) established Central and South America Cercosporella dolichandrae Crous & Leaf spot Not released, occurs Moderate den Breeӱen (Mycosphaerellales: pathogen locally; also under Mycosphaerellaceae) investigation

Charidotis auroguttata Boheman Leaf feeder Released 1999, Trivial Miranda, Venezuela Sparks 1999; Williams (Coleoptera: Chrysomelidae: established 2002 Cassidinae)

Hedwigiella jureceki (Obenberger) Leaf miner Released 2007, Trivial Misiones, Argentina King et al. 2011b (Coleoptera: Buprestidae) established

Hypocosmia pyrochroma Jones Leaf feeder Released 2010, Misiones, Argentina Dhileepan et al. 2007; (Lepidoptera: Pyralidae) establishment King et al. 2011b unconfirmed Tecoma stans (L.) Juss ex Kunth Under Mada polluta (Mulsant) (Coleoptera: Leaf feeder Released 2013, Moderate Guerrero, Mexico Madire et al. 2011a; var. stans evaluation Coccinellidae) established Madire 2013 yellow bells North and Central America Prospodium transformans (Ellis & Leaf rust Released 2010, not Mexico Wood 2008; Madire et al. Everh.) Cummins (Pucciniales: pathogen established 2011a Uropyxidaceae)

Pseudonapomyza sp. (Diptera: Leaf miner Released 2014, Trivial San Pedro, Madire 2009/10; Madire et Agromyzidae) established Argentina al. 2011a,b

CACTACEAE Austrocylindropuntia subulata Not Cactoblastis cactorum (Berg) Cladode borer Alternative host, Unknown Argentina via Moran & Zimmermann (Muehlenpf.) Backeb. (=Opuntia determined (Lepidoptera: Pyralidae) established Australia 1991b exaltata (Berger)) long spine cactus South America Cereus jamacaru DC. and C. Complete Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Extensive Argentina via Moran & Zimmermann hildmannianus K.Schum. Trelles) (Hemiptera: Pseudococcidae) established Australia 1991b; Klein 1999; queen of the night Paterson et al. 2011 South America Nealcidion cereicola (Fisher) [formerly Stem borer Alternative host, Considerable Argentina via Klein 1999 in Alcidion] (Coleoptera: released 1990, Australia Cerambycidae) established

Cylindropuntia fulgida (Engelm.) Complete Dactylopius tomentosus (Lamarck), Cladode sucker Released 2008, Extensive Mexico (Baja Mathenge et al. 2009a,b, F.M.Knuth var. fulgida Zim 'cholla' biotype (Hemiptera: established California) 2010a,b; Paterson et al. chain-fruit cholla Dactylopiidae) 2011

North and Central America Dactylopius tomentosus (Lamarck), Cladode sucker Alternative host, Trivial South-Western USA Moran & Zimmermann 'imbricata' biotype (Hemiptera: released 1970, via Australia 1991b Dactylopiidae) established

Cylindropuntia fulgida (Engelm.) Complete Dactylopius tomentosus (Lamarck), Cladode sucker Released 2008, Extensive Mexico (Baja F.M.Knuth var. mamillata (Engelm.) 'cholla' biotype (Hemiptera: established California) Backeb. Dactylopiidae) boxing-glove cactus North and Central America Cylindropuntia imbricata (Haw.) Substantial Dactylopius tomentosus (Lamarck), Cladode sucker Released 1970, Considerable South-Western USA Moran & Zimmermann F.M.Knuth 'imbricata' biotype) (Hemiptera: established via Australia 1991b; Mathenge et al. imbricate prickly pear Dactylopiidae) 2009a

North and Central America Metamasius spinolae (Gyllenhal) Stem borer Alternative host, Mexico Moran & Zimmermann (Coleoptera: Curculionidae) released 1974, not 1991b established Cylindropuntia leptocaulis (DC.) Complete Dactylopius tomentosus (Lamarck) Cladode sucker Alternative host, Extensive South-Western USA Moran & Zimmermann F.M.Knuth ('imbricata' biotype) (Hemiptera: released 1977, via Australia 1991b; Mathenge et al. pencil cactus Dactylopiidae) established 2009a

North and Central America Harrisia bonplandii (Parm.) Britton & Substantial Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Considerable Argentina via Rose (= H. balansae (K.Schum.) Trelles) (Hemiptera: Pseudococcidae) released 2006, Australia N.P.Taylor & Zappi) established angular cactus, strangler cactus South America Harrisia martinii (Labour.) Complete Hypogeococcus festerianus (Lizer y Stem sucker Released 1983, Extensive Argentina via Moran & Zimmermann moon cactus Trelles) (Hemiptera: Pseudococcidae) established Australia 1991b; Klein 1999

South America Nealcidion cereicola Fisher [formerly Stem borer Released 1990, Considerable Argentina via Klein 1999 in Alcidion] (Coleoptera: established Australia Cerambycidae) Harrisia pomanensis (F.A.C.Weber) Substantial Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Considerable Argentina via Britton & Rose Trelles) (Hemiptera: Pseudococcidae) released 2013, Australia midnight lady, devil's rope cactus established South America Harrisia tortuosa (J.Forbes ex Otto & Substantial Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Considerable Argentina via A.Dietr.) Britton & Rose Trelles) (Hemiptera: Pseudococcidae) released 2014, Australia spiny snake cactus established South America Hylocereus undatus (Haw.) Britton & Not Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Unknown Argentina via Rose determined Trelles) (Hemiptera: Pseudococcidae) established Australia dragon fruit Tropical America Opuntia aurantiaca Lindl. Substantial Cactoblastis cactorum (Berg) Cladode borer Alternative host, Moderate Argentina via Moran & Annecke 1979; jointed cactus (Lepidoptera: Pyralidae) established Australia Moran & Zimmermann 1991a; Zimmermann et al. South America 2004 Dactylopius austrinus De Lotto Cladode sucker Released 1935, Extensive Central & Western Moran & Annecke 1979; (Hemiptera: Dactylopiidae) established Argentina via Moran & Zimmermann Australia 1991a Mimorista pulchellalis Dyar Cladode borer Released 1979, not Argentina (Parana) Moran & Zimmermann (Lepidoptera: Crambidae) established 1991a

Nanaia sp. (Lepidoptera: Pyralidae) Cladode borer Released 1983, not Peru Moran & Zimmermann established 1991a

Zophodia tapiacola (Dyar) [formerly in Cladode borer Released 1976 & Argentina (Formosa Moran & Zimmermann Tucumania] (Lepidoptera: Pyralidae) 1982, not & Buenos Aires 1991a established Provinces) Opuntia engelmannii Salm-Dyck ex Negligible Cactoblastis cactorum (Berg) Cladode borer Alternative host, Trivial on large Argentina via Engelm. (=O. lindheimeri, O. to (Lepidoptera: Pyralidae) released 1938, plants, moderate Australia tardospina) substantial, established on small plants small round-leaved prickly pear depending on variety North and Central America Dactylopius opuntiae (Cockerell), Cladode sucker Alternative host, Moderate to trivial, USA (Texas, Moran & Zimmermann 'ficus' biotype (Hemiptera: released 1938, depending on Arizona) via 1991b Dactylopiidae) established climate & plant Australia variety Dactylopius opuntiae (Cockerell), Cladode sucker Alternative host, Extensive, in Via Australia 'stricta' biotype (Hemiptera: released 2012, combination with Dactylopiidae) established pathogen

Opuntia ficus-indica (L.) Mill. Substantial Cactoblastis cactorum (Berg) Cladode borer Released 1933, Extensive Argentina via Annecke & Moran 1978; mission prickly pear, sweet prickly (Lepidoptera: Pyralidae) established Australia Zimmermann & Moran pear 1991 Dactylopius opuntiae (Cockerell), Cladode sucker Released 1938, Extensive USA (Texas, Annecke & Moran 1978; Central America 'ficus' biotype (Hemiptera: established Arizona) via Zimmermann & Moran Dactylopiidae) Australia 1991

Lagocheirus funestus (Thompson) Stem borer Released 1943, Trivial Mexico via Australia Annecke & Moran 1978; [formerly in Archlagocheirus] established Zimmermann & Moran (Coleoptera: Cerambycidae) 1991

Metamasius spinolae (Gyllenhal) Stem borer Released 1948, Extensive Mexico Zimmermann & Moran (Coleoptera: Curculionidae) established 1991

Opuntia humifusa (Raf.) Raf. Complete Dactylopius opuntiae (Cockerell), Cladode sucker Alternative host, Considerable in Via Australia creeping prickly pear, large-flowered Australian ('stricta') biotype established hot, arid regions, prickly pear (Hemiptera: Dactylopiidae) moderate in cooler regions & Highveld North America Opuntia monacantha Haw. Complete Cactoblastis cactorum (Berg) Cladode borer Alternative host, Considerable Argentina via Moran & Zimmermann smooth/drooping prickly pear (Lepidoptera: Pyralidae) established Australia 1991b; Zimmermann et al. 2009 South America Dactylopius ceylonicus (Green) Cladode sucker Released 1913, Extensive Temperate South Moran & Zimmermann (Hemiptera: Dactylopiidae) established America, Brazil via 1991b; Zimmermann et al. Sri Lanka 2009 Opuntia salmiana J.Parm. ex Pfeiff. Negligible Cactoblastis cactorum (Berg) Cladode borer Alternative host, Trivial to moderate Argentina via Moran & Zimmermann bur cactus (Lepidoptera: Pyralidae) established Australia 1991b

South America Opuntia spinulifera Salm-Dyck Not Cactoblastis cactorum (Berg) Cladode borer Alternative host, Unknown Argentina via Moran & Zimmermann large round-leaved prickly pear determined (Lepidoptera: Pyralidae) established Australia 1991b

Central America Opuntia stricta (Haw.) Haw. Substantial Cactoblastis cactorum (Berg) Cladode borer Alternative host, Extensive Argentina via Hoffmann et al. 1999 Australian pest pear (Lepidoptera: Pyralidae) released 1980s, Australia established North America and Caribbean Dactylopius opuntiae (Cockerell), Cladode sucker Released 1997, Extensive North America via Hoffmann et al.1999; Australian ('stricta') biotype established Australia Paterson et al. 2011 (Hemiptera: Dactylopiidae)

Opuntia tomentosa Salm-Dyck Substantial Dactylopius opuntiae (Cockerell), Cladode sucker Alternative host, Extensive USA (Texas, velvet opuntia, velvet tree-pear but 'ficus' biotype (Hemiptera: released 1938, Arizona) via localised Dactylopiidae) established Australia Mexico and Guatemala Peniocereus serpentinus (Lag. & Not Hypogeococcus festerianus (Lizer y Stem sucker Alternative host, Unknown Argentina via Rodr.) N.P.Taylor determined Trelles) (Hemiptera: Pseudococcidae) established Australia serpent cactus Mexico Pereskia aculeata Mill. Under Catorhintha schaffneri Brailovsky & Stem wilter Released 2014, Unknown Santa Catarina, Paterson et al. 2014a,b Barbados gooseberry, pereskia evaluation Garcia (Hemiptera: Coreidae) established Brazil

South America and Caribbean Phenrica guerini Bechyné Leaf feeder Released 1991 & Extensive locally Rio de Janiero, Klein 1999; Paterson et al. (Coleoptera: Chrysomelidae: 1995, established Brazil 2011, 2012 Galerucinae)

CLUSIACEAE Hypericum perforatum L. Complete Agrilus (Spiragrilus) hyperici Root borer Released 1974- France via Australia Gordon & Kluge 1991 St John's wort (Creutzer) (Coleoptera: Buprestidae) 1979, not & California (USA) established Europe and Asia Aphis chloris Koch (Hemiptera: Sap sucker Released 1982, not Germany via Gordon & Kluge 1991 Aphididae) established Canada

Aplocera efformata (Guenée) Leaf feeder Released 1983, not France via Australia Gordon & Kluge 1991 (Lepidoptera: Geometridae) established

Chrysolina hyperici(Forster) Leaf feeder Released 1960- England via Gordon & Kluge 1991 (Coleoptera: Chrysomelidae: 1973, not Australia (1960); Chrysomelinae) established England via Australia via USA via Canada (1973) Chrysolina quadrigemina Suffrian Leaf feeder Released 1960- Extensive France via Australia Gordon & Kluge 1991 (Coleoptera: Chrysomelidae: 1973, established Chrysomelinae)

Zeuxidiplosis giardi (Kieffer) (Diptera: Shoot-tip galler Released 1972, Moderate France via California Gordon & Kluge 1991 Cecidomyiidae) established (USA) via Australia

COMMELINACEAE Tradescantia fluminensis Vell. Not Neolema abbreviata (Lacordaire) Shoot-tip and Release permit Brazil via New Fowler et al., 2013 wandering Jew, spiderwort applicable (Coleoptera: Chrysomelidae: leaf feeder issued Zealand Criocerinae) South America CONVOLVULACEAE Convolvulus arvensis L. Not Aceria malherbae Nuzzaci (Acari: Leaf galler Released 1994, not Greece Craemer 1995 field bindweed applicable Trombidiformes: Eriophyidae) established

Europe and Asia Septoria sp. (anamorphic fungus; Leaf spot Not released, occurs Trivial Mycosphaerellales: pathogen locally; rejected Mycosphaerellaceae)

FABACEAE Acacia baileyana F.Muell. Negligible Dasineura pilifera Kolesik (Diptera: Flower galler Released 2016, Unknown Victoria, Australia Bailey’s wattle Cecidomyiidae) established

Australia Melanterius maculatus Lea Seed feeder Alternative host, Moderate NSW, Australia Dennill et al. 1999; Impson (Coleoptera: Curculionidae) released 2006 & et al. 2009, 2011 2009, established Acacia cyclops A.Cunn. ex G.Don Substantial Calonectria scoparia Ribeiro & Leaf spot Not released, occurs Trivial red eye, rooikrans Matsuoka ex Peerally (Hypocreales: pathogen locally; rejected Nectriaceae) (anamorphic fungus: Australia Cylindrocladium candelabrum)

Dasineura dielsi Rübsaamen (Diptera: Flower galler Released 2002, Extensive WA, Australia Adair 2005; Impson et al. Cecidomyiidae) established 2009, 2011

Melanterius servulus Pascoe Seed feeder Alternative host, Considerable WA, Australia Dennill et al. 1999; Impson (Coleoptera: Curculionidae) released 1991 & et al. 2009, 2011 1994, established Pseudolagarobasidium acaciicola Die-back Not released, occurs Considerable Wood & Ginns 2006, Ginns (Polyporales: pathogen locally; under Kotzé et al. 2015 Phanerochaetaceae) investigation

Acacia dealbata Link Negligible Dasineura pilifera Kolesik (Diptera: Flower galler Released 2016, Unknown Victoria, Australia silver wattle Cecidomyiidae) established

Australia Melanterius maculatus Lea Seed feeder Alternative host, Moderate Australia Dennill et al. 1999; Impson (Coleoptera: Curculionidae) released 1994-2007, et al. 2009, 2011 established Acacia decurrens (Wendl.) Willd. Negligible Dasineura pilifera Kolesik (Diptera: Flower galler Release permit Victoria, Australia green wattle Cecidomyiidae) issued

Australia Melanterius maculatus Lea Seed feeder Alternative host, Moderate Victoria, Australia Dennill et al. 1999; Impson (Coleoptera: Curculionidae) released 2000-2007, et al. 2009, 2011 established Acacia longifolia (Andr.) Willd. Substantial Melanterius ventralis Lea (Coleoptera: Seed feeder Released 1985, Extensive Australia Dennill & Donnelly 1991; long-leaved wattle Curculionidae) established Dennill et al. 1999; Impson et al. 2009, 2011 Australia Trichilogaster acaciaelongifoliae Bud galler Released 1982, Extensive Australia Dennill & Donnelly 1991; (Froggatt) (Hymenoptera: established Dennill et al. 1999; Impson Pteromalidae) et al. 2009

Acacia mearnsii De Wild. Substantial Ceratocystis albofundus M.J.Wingf., Gummosis Not released, occurs Moderate, Morris et al. 1999 black wattle De Beer & M.J.Morris (Microascales: pathogen locally; shelved localised Ceratocystidaceae) Australia Cylindrobasidium laeve (Pers.) Early succession Not released, occurs Extensive when Morris et al. 1999 Chamuris (Polyporales: wood rot locally; developed applied according Physalacriaceae) pathogen, as cut stump to label prevents inoculant instructions coppice growth "Stumpout" Dasineura rubiformis Kolesik (Diptera: Flower galler Present before Extensive in WC, Australia Adair 2004; Impson et al. Cecidomyiidae) release, established Moderate EC, 2008, 2009, 2011 Trivial MP & KZN Melanterius maculatus Lea Seed feeder Released 1993- Moderate NSW, ACT, Dennill et al. 1999; Impson (Coleoptera: Curculionidae) 1996, established Australia et al. 2009, 2011

Acacia melanoxylon R.Br. Substantial Melanterius acaciae Lea (Coleoptera: Seed feeder Released 1986, Extensive Australia Dennill et al. 1999; Impson Australian blackwood Curculionidae) established et al. 2009, 2011

Australia Acacia podalyriifolia A.Cunn. ex Negligible Melanterius maculatus Lea Seed feeder Alternative host, Moderate but QLD, Australia Dennill et al. 1999; Impson G.Don. (Coleoptera: Curculionidae) released 2008, localised et al. 2009, 2011 pearl acacia established Australia

Acacia pycnantha Benth. Substantial Cylindrobasidium laeve (Pers.) Early succession Alternative host; not Extensive when Morris et al. 1999 golden wattle Chamuris (Polyporales: wood rot fungus released, occurs applied according Physalacriaceae) locally; developed to label Australia as cut stump instructions inoculant "Stumpout", but never used Melanterius maculatus Lea Seed feeder Alternative host, Moderate SA, Vic, Australia Dennill et al. 1999; Impson (Coleoptera: Curculionidae) released 2005-2007, et al. 2009, 2011 established Trichilogaster signiventris (Girault) Bud galler Released 1987 & Considerable/ Australia Hoffmann et al. 2002; (Hymenoptera: Pteromalidae) 1992, established Extensive Impson et al. 2009, 2011

Acacia saligna (Labill.) H.L.Wendl. Substantial Melanterius compactus Lea Seed feeder Released 2001 & Considerable WA, Australia Dennill et al. 1999; Impson Port Jackson willow (Coleoptera: Curculionidae) 2003, established et al. 2009, 2011

Australia Uromycladium morrisii Doungsa-ard, Gall former Released 1987, Extensive WA, Australia Morris 1987, 1991, 1997, McTaggart, Geering & R.G. Shivas established 1999; Wood & Morris 2007 (Pucciniales: Pileolariaceae)

Caesalpinia decapetala (Roth) Negligible Sulcobruchus subsuturalis (Pic) Seed feeder Released 1999, Trivial India Coetzer & Neser 1999; Alston (Coleoptera: Chrysomelidae: established Coetzer 2000; Byrne et al. Mauritius thorn Bruchinae) 2011

Asia Gleditsia triacanthos L. Not Megabruchidius tonkineus (Pic) Seed feeder Not released, occurs Unknown honey locust determined (Coleoptera: Chrysomelidae: locally Bruchinae) North America Leucaena leucocephala (Lam.) de Negligible Acanthoscelides macrophthalmus Seed feeder Released 1999, Trivial Mexico Shoba & Olckers 2010; Wit (Schaeffer) (Coeloptera: established Olckers 2011a; Sharratt & leucaena Chrysomelidae: Bruchinae) Olckers 2012

Tropical America Paraserianthes lophantha (Willd.) Substantial Melanterius servulus Pascoe Seed feeder Released 1989, Considerable Australia Dennill et al. 1999; Impson Nielsen (formerly Albizia lophantha (Coleoptera: Curculionidae) established et al. 2009, 2011 Willd.) stink bean Uromycladium woodii Doungsa-ard, Gall former Released 2016, Unknown WA, Australia Australia McTaggart, Geering & R.G. Shivas established (Pucciniales: Pileolariaceae)

Prosopis L. species Negligible Algarobius bottimeri Kingsolver Seed feeder Released 1990, not South-western USA Impson et al. 1999 mesquite, prosopis (Coleoptera: Chrysomelidae: established Bruchinae) North and Central America Algarobius prosopis (LeConte) Seed feeder Released 1987, Considerable Arizona, USA Hoffmann et al.1993; (Coleoptera: Chrysomelidae: established Impson et al. 1999; Bruchinae) Roberts 2006; Van Klinken et al. 2009; Zachariades et al. 2011a Neltumius arizonensis (Schaeffer) Seed feeder Released 1993, Unknown Arizona, USA Coetzer & Hoffmann 1997; (Coleoptera: Chrysomelidae: established Impson et al. 1999; Bruchinae) Strathie 1995

Sesbania punicea (Cav.) Benth. Complete Neodiplogrammus quadrivittatus Stem borer Released 1984, Extensive Argentina (Buenos Hoffmann & Moran 1999 red sesbania (Olivier) (Coleoptera: Curculionidae) established Aires Prov); Brazil (Rio Grande do Sul) South America Rhyssomatus marginatus Fåhraeus Seed feeder Released 1984, Extensive Argentina (Buenos Hoffmann & Moran 1999 (Coleoptera: Curculionidae) established Aires Prov)

Trichapion lativentre (Béguin- Flower-bud Released 1970s, Extensive Argentina (Buenos Hoffmann & Moran 1999 Billecocq) (Coleoptera: Brentidae: feeder established, present Aires Prov) Apioninae) before release

HALORAGACEAE Myriophyllum aquaticum (Vell.) Complete Lysathia sp. (Coleoptera: Leaf feeder Released 1994, Extensive Brazil Cilliers 1999; Coetzee et Verdc. Chrysomelidae: Galerucinae) established al. 2011b parrot's feather Xanthomonas campestris (Pammel) Bacterial wilt Not released, occurs Considerable, Morris et al. 1999 South America Dawson (?) (Proteobacteria: locally; rejected localised, not Xanthomonadales: persistent Xanthomonadaceae)

HYDROCHARITACEAE Hydrilla verticillata (L.f.) Royle Substantial Hydrellia purcelli Deeming (Diptera: Leaf miner Release permit Singapore Coetzee et al. 2011a; hydrilla Ephydridae) issued, shelved Bownes 2014, 2015; Bownes & Deeming 2016 Old World Parapoynx diminutalis Snellen Leaf cutter Not released, occurs Extensive Asia Bownes, 2010, 2018 (Lepidoptera: Crambidae) locally

MYRTACEAE Leptospermum laevigatum (Gaertn.) Negligible Aristaea thalassias (Meyrick) [formerly Leaf feeder Released 1996, Trivial Australia Gordon 1999b, Gordon F.Muell. in Parectopa] (Lepidoptera: established 2011 Australian myrtle Gracillariidae)

Australia Dasineura strobila Dorchin (Diptera: Bud galler Released 1994, Trivial Australia Gordon 1999b; Dorchin & Cecidomyiidae) established, present Adair 2011; Gordon 2011 before release POACEAE Arundo donax L. Negligible Tetramesa romana Walker Stem galler Present without Trivial Marshall et al. 2018 giant reed (Hymenoptera: Eurytomidae) - South being released African biotype Mediterranean, Europe and Asia PONTEDERIACEAE Eichhornia crassipes (C.Mart.) Substantial Acremonium zonatum (Sawada) Leaf pathogen Not released, occurs Trivial but can be Morris et al. 1999 Solms W.Gams (anamorphic fungus; locally considerable in water hyacinth Hypocreales) presence of agents South America Alternaria eichhorniae Nag-Raj & Leaf pathogen Not released, occurs Trivial but can be Morris et al. 1999 Ponnappa (anamorphic fungus; locally; shelved considerable in Pleosporales: Pleosporaceae) presence of insect agents Cercospora piaropi Tharp. Leaf pathogen Not released, occurs Unknown, cannot Morris et al. 1999 (anamorphic fungus; locally be distinguished Mycosphaerellales: from C. rodmanii Mycosphaerellaceae) in the field

Cercospora rodmanii Conway Leaf pathogen Released 1992, Considerable USA Hill & Cilliers 1999; Morris (anamorphic fungus; established et al. 1999 Mycosphaerellales: Mycosphaerellaceae)

Cornops aquaticum (Brüner) Leaf feeder Released 2011, Amazonas, Brazil Hill & Cilliers 1999; (Orthoptera: Acrididae: Leptysminae) establishment Bownes 2009, Bownes et unconfirmed al. 2010a,b; Coetzee et al. 2011b; Bownes et al. 2013a,b Eccritotarsus catarinensis (Carvalho) Leaf sucker Released 1996, Considerable Santa Catarina, Hill & Cilliers 1999; (Hemiptera: Miridae) established Brazil; and Loreto, Coetzee et al. 2005, 2007, Peru 2011b Eccritotarsus eichhorniae Henry Leaf sucker Released 2007, Considerable Loreto, Peru Paterson et al. 2016; (Hemiptera: Miridae) established Henry 2017

Megamelus scutellaris Berg Leaf sucker Released 2013, Unknown Argentina via USA Coetzee et al. 2011b (Hemiptera: Delphacidae) established

Neochetina bruchi Hustache Stem borer Released 1990 & Considerable Argentina via Hill & Cilliers 1999; (Coleoptera: Curculionidae) 1996, established Zimbabwe Coetzee et al. 2011b

Neochetina eichhorniae Warner Stem borer Released 1974 & Considerable Argentina Hill & Cilliers 1999; (Coleoptera: Curculionidae) 1985, established Coetzee et al. 2011b

Niphograpta albiguttalis Warren Petiole borer Released 1990, Considerable Argentina via USA Hill & Cilliers 1999; (Lepidoptera: Crambidae) established Coetzee et al. 2011b

Orthogalumna terebrantis Wallwork Leaf miner Released 1989, Considerable Argentina Hill & Cilliers 1999; (Acari: Sarcoptiformes: Galumnidae) established Coetzee et al. 2011b

PROTEACEAE Hakea gibbosa (Sm.) Cav. Negligible Aphanasium australe (Boisduval) Stem borer Alternative host, Unknown Australia (NSW) Gordon 1999a, 2003; rock hakea (Coleoptera: Cerambycidae) released 2001, Gordon & Fourie 2011 established Australia Erytenna consputa Pascoe Green-fruit Alternative host, Trivial Australia (SA) Gordon 1999a; Gordon & (Coleoptera: Curculionidae) feeder released 1975, Fourie 2011 established Hakea sericea Schrad. & J.C.Wendl. Substantial Aphanasium australe (Boisduval) Stem borer Released 2001, Unknown Australia (NSW) Gordon 1999a, 2003; silky hakea (Coleoptera: Cerambycidae) established Gordon & Fourie 2011

Australia Carposina autologa Meyrick Seed feeder Released 1970 & Trivial Australia (South Gordon 1999a; Gordon & (Lepidoptera: Carposinidae) 1982, established East & South West) Fourie 2011

Colletotrichum acutatum J.H. Stem gummosis Not released, occurs Extensive South Africa Morris 1991; Morris et al. Simmonds (Glomerellales: disease locally; developed 1999; Lubbe et al. 2004; Glomerellaceae) as mycoherbicide Gordon & Fourie 2011 "Hakatak" Cydmaea binotata Lea (Coleoptera: Leaf & shoot Released 1979, Trivial Australia Gordon 1999a; Gordon & Curculionidae) borer established Fourie 2011

Dicomada rufa Blackburn (Coleoptera: Flower-bud Released 2006, Unknown Australia Gordon 1999a; Gordon & Curculionidae) feeder established Fourie 2011

Erytenna consputa Pascoe Green-seed Released 1970 & Extensive Australia (SA) Gordon 1999a; Gordon & (Coleoptera: Curculionidae) feeder 1974, established Fourie 2011

SALVINIACEAE Salvinia molesta D.S.Mitch. Complete Cyrtobagous salviniae Calder & Stem borer Released 1985, Extensive Brazil via Australia Cilliers et al. 2003; water fern Sands (Coleoptera: Curculionidae) established via Namibia Coetzee et al. 2011b

South America SAPINDACEAE Cardiospermum grandiflorum Sw. Under Cissoanthonomus tuberculipennis Seed feeder Released 2013, Extensive Argentina Mc Kay et al. 2010; balloon vine evaluation Hustache (Coleoptera: Curculionidae) established Simelane et al. 2011a

South America SOLANACEAE Solanum elaeagnifolium Cav Substantial Frumenta nephelomicta (Meyrick) Fruit galler Released 1979 & Mexico Olckers et al. 1999 silverleaf nightshade, satansbos (Lepidoptera: Gelechiidae) 1985, not established North, Central and South America Frumenta sp. (Lepidoptera: Stem & fruit Released 1989, not Texas, USA Olckers 1995; Adamski & Gelechiidae) galler established Brown 2002

Leptinotarsa defecta (Stål) Leaf feeder Released 1992, Moderate Texas, USA Olckers & Hulley 1994; (Coleoptera: Chrysomelidae: established Olckers et al. 1995; Chrysomelinae) Hoffmann et al. 1998

Leptinotarsa texana (Schaeffer) Leaf feeder Released 1992, Extensive Texas, USA Olckers & Hulley 1994; (Coleoptera: Chrysomelidae: established Olckers et al. 1995; Chrysomelinae) Hoffmann et al. 1998

Solanum mauritianum Scop. Negligible Anthonomus santacruzi Hustache Flower-bud Released 2008, Trivial Argentina Olckers 2003, 2008, bugweed (Coleoptera: Curculionidae) feeder established 2011b; Cowie et al. 2016a, 2017; Singh & Olckers South America 2017; Cowie et al. 2018; English & Olckers 2018 Gargaphia decoris Drake (Hemiptera: Leaf sucker Released 1999, Trivial, occasional Argentina, Brazil Olckers 2000a, 2011b; Tingidae) established outbreaks Olckers & Lotter 2004; Olckers & Borea 2009; Cowie et al. 2016b, 2018 Solanum sisymbriifolium Lam. Substantial Gratiana spadicea (Klug) (Coleoptera: Leaf feeder Released 1994, Extensive Argentina, southern Hill & Hulley 1995; Byrne wild tomato, dense-thorned bitter Chrysomelidae: Cassidinae) established Paraguay and Brazil et al. 2002; King et al. apple 2011a South America VERBENACEAE Lantana camara L. (sensu lato) Negligible Aceria lantanae (Cook) (Acari: Flower galler Released 2007, Extensive, on Florida, USA; Cuba Urban et al. 2001, 2011a; lantana (Highveld) Trombidiformes: Eriophyidae) established certain varieties in Mukwevho et al. 2017 to some areas Central and South America substantial Alagoasa parana Samuelson Leaf feeder Released 1985, not Brazil via Australia Cilliers & Neser 1991 (coastal & (Coleoptera: Chrysomelidae: established Lowveld) Galerucinae)

Autoplusia illustrata (Guenée) Leaf feeder Released 1978 & Colombia via Cilliers & Neser 1991 (Lepidoptera: Noctuidae) 1984, not Australia established Calycomyza lantanae (Frick) (Diptera: Leaf miner Released 1982, Moderate Trinidad via Baars & Neser 1999 Agromyzidae) established Australia; Later also Florida, USA Coelocephalapion camarae Kissinger Petiole galler Released 2007, Moderate but Mexico Baars et al. 2006 (Coleoptera: Brentidae: Apioninae) established localized

Crocidosema lantana Busck [formerly Flower & Present before Trivial Mexico via Hawaii, Baars 2003 in Epinotia] (Lepidoptera: Tortricidae) receptacle miner release; released USA 1984, established Eutreta xanthochaeta Aldrich (Diptera: Shoot galler Released 1983, not Hawaii, USA Mabuda 2005 Tephritidae) established; rejected

Falconia intermedia (Distant) Leaf sucker Released 1999, Moderate, Jamaica Baars et al. 2003 (Hemiptera: Miridae) established localized

Hypena laceratalis Walker Leaf feeder Not released, native Moderate Native Baars 2003 (Lepidoptera: Noctuidae)

Lantanophaga pusillidactyla (Walker) Flower miner Present before Trivial Mexico via Hawaii, Baars 2003 (Lepidoptera: Pterophoridae) release; released USA 1984, not established Leptobyrsa decora Drake (Hemiptera: Leaf sucker Released 1972- Colombia, Peru via Cilliers & Neser 1991 Tingidae) 1984, not Australia established Longitarsus bethae Savini & Escalona Root feeder Released 2007, Moderate, Mexico Simelane 2005, 2010 (Coleoptera: Chrysomelidae: established localised Galerucinae)

Neogalia sunia (Guenée) Leaf feeder Released 1962 & California, USA via Cilliers & Neser 1991 (Lepidoptera: Noctuidae) 1969, not Hawaii via Australia established Octotoma championi Baly Leaf miner Released 1978, Costa Rica via Cilliers & Neser 1991 (Coleoptera: Chrysomelidae: 1995, not Australia Cassidinae) established

Octotoma scabripennis Guèrin- Leaf miner Released 1971- Considerable Mexico, via Hawaii, Cilliers 1987 Mèneville (Coleoptera: 1975, established via Australia Chrysomelidae: Cassidinae)

Ophiomyia camarae Spencer Leaf miner Released 2001, Considerable Florida, USA Simelane 2002; Simelane (Diptera: Agromyzidae) established & Phenye 2005

Ophiomyia lantanae (Froggatt) Fruit miner Present before Moderate Mexico via Hawaii, Cilliers 1987 (Diptera: Agromyzidae) release; released USA 1961 Orthonama ignifera (Warren) [formerly Leaf feeder Release permit Florida, USA; Williams & Madire 2008 in Leptostales] (Lepidoptera: issued Mexico Geometridae)

Passalora lantanae (Chupp) U.Braun Leaf pathogen Released 2002, not Florida, USA Den Breeÿen 2003; Den & Crous var. lantanae [formerly in established Breeÿen & Morris 2003; Mycovellosiella] (anamorphic fungus; Retief 2010a Mycosphaerellales: Mycospheraellaceae)

Plagiohammus spinipennis (Thomson) Stem borer Released 1973, not Mexico via Hawaii, Cilliers & Neser 1991 (Coleoptera: Cerambycidae) established USA via Australia

Puccinia lantanae Farl. (Pucciniales: Leaf and stem Release permit Peru Renteria et al. 2004 Pucciniaceae) rust pathogen issued; imported

Salbia haemorrhoidalis (Guenée) Leaf feeder Released 1962, Trivial Cuba via Hawaii, Baars 2003 (Lepidoptera: Crambidae) established USA

Teleonemia elata Drake (Hemiptera: Leaf sucker Released 1972, not Brazil via Australia Cilliers & Neser 1991 Tingidae) established

Teleonemia scrupulosa Stål Leaf & flower Released 1961, Considerable Mexico via Hawaii Baars & Neser 1999 (Hemiptera: Tingidae) sucker 1972, 1984, (1961); C & S established America via Australia (1972); via Mauritius (1984) Uroplata fulvopustulata Baly Leaf miner Released 1978, not Costa Rica via Day et al. 2003 (Coleoptera: Chrysomelidae: established Australia Cassidinae)

Uroplata girardi Pic (Coleoptera: Leaf miner Released 1974, Considerable, Brazil via Hawaii, Baars & Neser 1999 Chrysomelidae: Cassidinae) 1983, established coastal USA via Australia (1974); Argentina via Australia (1983) Uroplata lantanae Buzzi & Winder Leaf miner Released 1984, not Brazil via Australia Cilliers & Neser 1991 (Coleoptera: Chrysomelidae: established Cassidinae)

For each of the agents that has established successfully, a rating is given for the perceived degree of damage that it inflicts on the target weed. These ratings (Anon. 1999), which have been widely adopted since 1999, and slightly amended in this account, are sometimes based on formal measurements and otherwise on subjective visual assessments, and include the following categories. Extensive: Very high levels of damage, as much as could be expected from the agent. Few plants survive, or growth is arrested, or almost no seeds are produced. Considerable: High levels of damage. Some plants may survive but growth rates are noticeably slower, or seed production is reduced by more than 50%. Moderate: Perceivable damage, but most plants survive. Growth may be slowed to some extent, or seed production is reduced by less than 50%. Trivial: Some damage, but survival, growth and seed production of the plants is almost normal. Unknown: Agent has been too recently released, or has not been evaluated.

For each weed species on which at least one biological control agent has established, the impact of the agent (where only one is established) or combined impact of all agents (where two are more are established) (i.e. the degree of control) (adapted from Anon. 1999) is rated by estimating the degree to which the impact or importance of the target weed species has been reduced by the agent(s). Assessments are based on the degree of reduction in the use of alternative control methods (chemical or mechanical) since the introduction of biological control agents, and include the following categories. Complete: No other control measures are needed to reduce the weed to acceptable levels, at least in areas where the agents are established. Substantial: Other methods are needed to reduce the weed to acceptable levels, but less effort is required (e.g. less frequent herbicide applications or less herbicide needed per unit area). Negligible: In spite of damage inflicted by the agents, control of the weed remains entirely reliant on the implementation of other control measures. Not determined: Either the release of the agents has been too recent for meaningful evaluation, or the programme has not been evaluated. Under evaluation: an active programme is underway to evaluate the impact of biological control agents on the weed, which is expected to result in categorisation as complete, substantial or negligible control.

Adapted from: Klein, H. 2011. A catalogue of the insects, mites and pathogens that have been used or rejected, or are under consideration, for the biological control of invasive alien plants in South Africa. In: Moran, V.C., Hoffmann, J.H. & Hill, M.P. (Eds) Biological Control of Invasive Alien Plants in South Africa (1999-2010). African Entomology (Special Issue) 19(2): 515-549.

Last updated July 2018. Contact: Costas Zachariades at e-mail: [email protected]