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230 Session 5 Prospects for Weed Biological Control in Pacific Islands

Releases of Natural Enemies in since 1980 for Classical Biological Control of Weeds

P. Conant1, J. N. Garcia2, M. T. Johnson3, W. T. Nagamine2, C. K. Hirayama1, G. P. Markin4 and R. L. Hill5

1Hawaii Department of Agriculture, Pest Control Branch, Hilo, Hawaii 96720 USA 2Hawaii Department of Agriculture, Plant Pest Control Branch, Honolulu, Hawaii 96814 USA 3USDAForest Service, Pacific Southwest Research Station, Institute of Pacific Islands Forestry, Volcano, Hawaii 96785 USA 4USDAForest Service, Bozeman, Montana 59717 USA, Retired 5Richard Hill and Associates, Private Bag 4704, Christchurch,

Abstract

A comprehensive review of biological control of weeds in Hawaii was last published in 1992, covering 74 natural enemy released from 1902 through 1980. The present review summarizes releases of 21 natural enemies targeting seven invasive weeds from 1981 to 2010. These projects were carried out by Hawaii Department of Agriculture (HDOA), USDA Forest Service (USFS), University of Hawaii (UH), and US Geological Survey Biological Resources Discipline. An appendix summarizing the chronology and outcomes of releases is included (Appendix 1).

Introduction (1998). Hawaii’s weed biological control projects from 1902 to1980 resulted in establishment of 42 and one fungal agent on 19 target weeds. The practice of classical biological control of Our objective here is to provide an updated weeds began in Hawaii in 1902, with the release report of agents released for weed biological control by the Territory of Hawaii of a tingid lacebug in Hawaii. We briefly summarize results for each (Teleonemia scrupulosa Stål) for control of lantana natural enemy released since 1981, and we include (Lantana camara L.) (Swezey, 1924). Since that time, an appendix listing all weed biological control agents Hawaii has witnessed several spectacular successes released in Hawaii from 1902 to 2010 (Appendix 1). in weed biological control, and today continues the

science of managing weeds at the landscape level using natural enemies introduced from the target’s Methods native range after thorough testing and evaluation. The most recent comprehensive review of weed Data on weed targets and weed biological biological control in Hawaii (Markin et al., 1992) control agents released since 1981 were compiled summarized 74 introductions of natural enemies from published and unpublished sources, including between 1902 and 1980. Additional information for records of the Hawaii Department of Agriculture introductions through the mid-1990s was included and personal observations of biological control in the worldwide compilation by Julien and Griffiths specialists and weed management partners.

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Results known to be active in the study area and could utilize P. perelegans in lab tests, no field-collected larvae or pupae were found to be parasitized (Ramadan et al., tarminiana Coppens & V. E. Bar- 2008). Trichogramma chilonis (Trichogrammatidae) ney (), banana poka parasitism affected from 0 to 26% of field collected eggs (Ramadan et al., 2008), and may be a significant Passiflora tarminiana (formerly P. mollissima) factor suppressing P. perelegans (Campbell et al., is a native to the South American , 1995). Several common predators of lepidopteran apparently introduced into Hawaii in the early 1900s larvae were noted also, but definitive evidence (Wagner et al., 1990). It is invasive in native forests of their impacts on P. perelegans was lacking on the islands of Hawaii and Kauai, and is found in (Campbell et al., 1995; Ramadan et al., 2008). alien forest habitat in the Kula area of Maui (Hauff, 2006). Funding for the biological control program passiflorae Syd. (Mycosphaerellaceae) in Hawaii came primarily from the USFS and Septoria passiflorae, a originally from National Park Service. P. tarminiana also is invasive Colombia, attacks P. tarminiana leaves, first forming in New Zealand, where a biological control program distinct yellow spots which eventually spread is being implemented (Williams and Hayes, 2007). to cover much of the leaf and cause premature abscission. It was released on Hawaii Island in 1996 Scea (= Cyanotricha) necyria (Felder & Rogen- (Trujillo et al., 2001), and quickly controlled a large hofer) () infestation of banana poka, which had smothered Scea necyria was imported from Colombia and native forest canopy at approximately 2,000 m subsequently released on the islands of Hawaii, Kauai elevation. S. passiflorae was much less effective in and Maui, but failed to establish. Despite repeated the drier habitat of Kula, Maui (G. Shishido, pers. releases between 1988 and 1992 totaling over 15,000 comm.). In Kokee, Kauai, defoliation of was individuals (including adult , defoliating observed at inoculation sites, but the clumped larvae, and eggs), no evidence of establishment was distribution of the weed in the forest may have ever observed. Larvae and pupae collected from the inhibited dispersal of the pathogen (G. Kawakami, field exhibited 17% and 9% parasitism, respectively, pers. comm.). Furthermore, the climate in Kokee is while no parasites were recovered from field collected somewhat drier than the windward inoculation sites eggs (Campbell et al., 1995). It was suspected that on the island of Hawaii, where infection and control S. necyria pupae may have been significantly preyed results were substantial (Trujillo, 2005). upon by birds. Research in Colombia has suggested that adult moths may be missing an obligate Coccinia grandis (L.) Voigt (Cucurbitaceae), nectar source in the Hawaii environments where ivy gourd this species was released (Campbell et al., 1995). Ivy gourd is probably native to central East Africa, Pyrausta perelegans Hampson (Crambidae) and was most likely moved to the Indo-Malayan Pyrausta perelegans imported from region in centuries past. It may have come to Hawaii was released on Hawaii, Maui, and Kauai islands as a food or medicinal plant, via immigrants from in 1991-1992, but established only on Hawaii and that region. Ivy gourd was first reported on Oahu Maui. Larvae feed inside flower buds, consuming in 1968, and has since spread to Hawaii, Maui, the ovary, anther, gynophore, inner flower tube, Kauai and Lanai. This climbing vine blanketed and (Ramadan et al., 2008). P. perelegans large areas of alien wayside trees and shrubs on populations have remained at low levels, with vine the island of Oahu, and less so in drier Kona infestation rates averaging 2-11% in post-release (Hawaii). Three insects were released for biological monitoring in 1992-1993 by Ramadan et al. (2008), control between 1996 and 1999 (Chun, 2002). and around 2% or less of flower buds infested a decade later (M.T. Johnson, unpublished data). Melittia oedipus Oberthür (Sessiidae) While generalist lepidopteran parasitoids were Melittia oedipus, a clear-winged from

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Kenya, was released from 1996 through 2002. mechanical control on the ground. On Hawaii, Larvae feed in both roots and stems of the vine, the task is even more daunting, given the large area forming galls. Galls can grow larger than 1.5 cm invaded, and resources are presently not available to and cause breakage of the vine, so that the foliage maintain extensive control. A single fungal pathogen above in the tree canopy dies. M. oedipus appears has been released for miconia biological control, in to be responsible for a widespread reduction in ivy both Hawaii and Tahiti. Additional agents are under gourd foliage density in tree and shrub canopies development (Johnson, 2010). on Oahu. Development of land for housing also contributed to the decline of ivy gourd in Colletotrichum gloeosporioides (Penz.) f. sp. the Kona area on Hawaii Island. Rat predation miconiae Killgore et al. (Glomerellaceae) on M. oedipus larvae and pupae appeared to be This anthracnose fungus, released in 1997, significant, but the potential for biotic interference is currently the only natural enemy approved for has not been studied further (Chun, 2002). miconia control in Hawaii (Killgore, 2002). Although laboratory tests by Meyer et al. (2008) found Acythopeus coccineae O’Brien and Pakaluk, significant mortality of miconia seedlings infected Acythopeus burkhartorum O’Brien and Pakaluk with C. g. f. sp. miconiae in Tahiti, the effects of the (Curculionidae) fungus on flowering and fruiting remain unknown. Two weevil species of the Acythopeus Post-release evaluation studies found this agent were first released in 1999. A. burkhartorum, a stem causing premature leaf drop in wild sapling gall former, failed to establish, despite many releases in Hawaii (Brenner, 2000). At higher elevations in different habitats. A. coccineae, whose larvae are in Tahiti, partial defoliation of large monospecific leaf miners, established on Oahu and in Kona, and stands have increased light levels penetrating appears capable of some impact. Ivy gourd has been the canopy so that some limited regeneration a target of physical containment on Kauai, Lanai and of native flora is occurring (Meyer et al., 2011). Maui, and these efforts, on Maui in particular, likely contributed to lack of establishment of A. coccineae Lantana camara L. (Verbenaceae), lantana there. The parasitoid Eupelmus prob. cushmani (Eupelmidae) has been reared from A. cocciniae larvae Lantana has been a target of weed biological and pupae, and may contribute to the low numbers of control work worldwide since 1902, when the A. cocciniae in field populations (M. Ramadan, pers. first ever natural enemy was released here in comm.). Both Acythopeus species were also released Hawaii (Swezey, 1924). A total of 25 agents for the biological control of ivy gourd in , have been released to control this weed in however, biotic interference by an unidentified Hawaii, with only one release made after 1980. hymenopteran parasitoid emerging from A. coccineae pupae was noted (Muniappan et al., 2009). Septoria sp. (Mycosphaerellaceae) Septoria sp. collected from lantana in Ecuador Miconia calvescens DC. (Melastomataceae), was first released in 1997 by Dr. Eduardo E. miconia Trujillo (UH) on the islands of Hawaii and Kauai (Trujillo, 2005). He reported excellent control Miconia was introduced to Hawaii as an of lantana at inoculated sites at Kokee on Kauai. ornamental tree around 1961 and planted on four of There has been no further update published on the major Hawaiian Islands before it was recognized the impact of the fungal pathogen on its host. as a serious invasive threat to native forests and watersheds (Conant et al., 1997). Miconia (L.) D. Don containment programs have been in effect since (Melastomataceae), Koster’s curse 1991 on Hawaii and Maui, and eradication programs continue on Oahu and Kauai. The containment Clidemia hirta was first collected in Hawaii in effort on Maui has been very expensive, entailing 1949, but was known to occur on Oahu since 1941 aerial application of and chemical/ (Wagner et al., 1990). It is now highly invasive on all

XIII International Symposium on Biological Control of Weeds - 2011 Session 5 Prospects for Weed Biological Control in Pacific Islands 233 the islands except Niihau and Kahoolawe, which are pteromalid wasp parasitoid has been reared from probably too dry. It is a large shrub that dominates the young fruit of Koster’s curse, suggesting possible the subcanopy of both native and alien wet biotic interference (T. Johnson, unpublished data). forests, and also can be a serious pest of windward pastures. Effectiveness of biological control agents Colletotrichum gloeosporioides f. sp. clidemiae released before 1981 is thought to be limited to Trujillo (Glomerellaceae) open, sunny habitats (Reimer and Beardsley, This pathogen was imported by Trujillo (2005) 1986; Reimer and Beardsley, 1988). As a result of from and released in 1986. Repeated HDOA exploration and testing in the Republic of annual inoculations were observed to reduce C. Trinidad and Tobago, four more agents were hirta cover in one area, and similar mycoherbicidal released between 1988 and 1995. Additional natural application has been pursued elsewhere on a enemies are still needed for sufficient control. limited basis. More recently, natural defoliation events affecting as much as 90% of C. hirta cover Lius poseidon Napp (Buprestidae) were observed on the wet windward side of Larvae of Lius poseidon, released in 1988, are Kauai, but plants appeared to recover with time leaf miners of C. hirta, while adult beetles are leaf (N. Barca, pers. comm.). This phenomenon has feeders (Conant, 2002). Larvae are parasitized been commonly observed on several islands at by at least one natural enemy, intentionally unpredictable intervals once the fungus became released for the control of agromyzid leaf miners well established in the 1990s. Cool, rainy, windy (Conant, 2002). Combined foliar damage by weather conditions appear conducive to outbreaks. adults and larvae to wild plants has been minimal. Morella (= Myrica) faya (Aiton) Wilbur Antiblemma acclinalis Hübner (Myricaceae), firetree (Noctuidae) Antiblemma acclinalis was first released in 1995 (Conant, 2009). This leaf feeder is recorded as Morella faya is thought to have been introduced established on Oahu and Kauai, but appears to never by Portuguese plantation workers who emigrated have become common, perhaps due to parasitism. from Atlantic islands in the 1800s. Firetree was used for reforestation in Hawaii in the 1920s, and bullata Meyrick now occurs on the islands of Hawaii, Maui, Lanai, () Oahu and Kauai (Whiteaker and Gardner, 1992). M. Carposina bullata, first liberated in 1995 failed faya is considered one of the worst to establish, possibly due to the low number of pupae in Hawaii Volcanoes National Park, since its ability that survived shipment from Tobago (Conant, 2009). to fix nitrogen allows it to thrive in low-nutrient lava Poor survival resulted in only about 140 adults being soils (Whiteaker and Gardner, 1985). A tortricid released, primarily on Hawaii and very few on Oahu. moth released for biological control of M. faya by HDOA in 1956 is considered ineffective, because it Mompha trithalama Meyrick only affected a minor population of Morella cerifera (Coleophoridae) (L.) Small (Markin, 2001). The National Park Service Mompha trithalama survived shipment from provided most of the funding for M. faya biological Tobago much better than C. bullata, resulting in a control efforts in the 1980s and early 1990s. Two higher number of individuals released on Hawaii natural enemies were released since 1980, but and Oahu in 1995 (Conant, 2009). M. trithalama neither appears to have any noticeable impact (G. was subsequently moved to Kauai and Maui. Markin, pers. observation). Larvae can now be found statewide, commonly feeding on the seeds of green fruit. It is possible Caloptilia coruscans (Walsingham) [= C. sp. that M. trithalama damage can result in premature nr. schinella (Walsingham)] (Gracillariidae) fruit drop, but the overall impact of this agent on Caloptilia coruscans, whose larvae are blotch seed production has not yet been quantified. A miners and leaf edge rollers, was released in 1991 from

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collections made in the Azores and (Markin, flowering (Markin, unpublished data). Three 2001). Populations have remained low, and foliar species of parasitoids attack A. ulicetella pupae, but damage has been minimal. Field collected larvae no larval parasitoids had ever been found, even were infected by a fungal pathogen, hymenopteran with intensive searching, and egg parasitism by parasitoids emerged from pupae, and lab tests found Trichogramma sp. appeared low (Markin et al., 1996). that local parasitoids attack its egg stage (Markin, 2001). Evidence of biotic interference also includes Stenopterapion scutellare (Kirby) (Brentidae) high levels of predation of C.coruscans larvae within leaf rolls (P. Yang and D. Foote, unpublished data). Stenopterapion scutellare was first liberated in 1961 (Markin and Yoshioka, 1989) on Maui Septoria hodgseii Gardner (186 released), but failed to establish and so was (Mycosphaerellaceae) re-released in 1989 on the island of Hawaii (788 Septoria hodgseii, a fungus collected from wax released). None of the characteristic galls caused myrtle, Morella cerifera, in North Carolina, was by the weevil were ever seen outside the field release tested for host specificity at the HDOA Plant cages in the 1990s, nor have they been seen since then. Pathogen Containment Facility in Honolulu and was subsequently released and briefly Sericothrips staphylinus Haliday (Thripidae), monitored on Hawaii Island in 1997 (Culliney et gorse thrips al., 2003). S. hodgsei was found established, but Two biotypes of this thrips, both a winged was never abundant in the experimental area, and brachypterous form, were released in Hawaii and no follow up work has been done. Smith between 1991 and 1994 (Markin et al., 1996). The (2002) mentions a personal communication from thrips feed on the mesophyll of gorse foliage, Trujillo, which suggested that acid rain near the producing pale stippled areas on leaves, spines, volcano could be inhibiting spore germination, and stems (Hill et al., 2001). Both biotypes became and that the fungus should be released elsewhere. established and widespread in the gorse infestation on Hawaii Island, but finding visible feeding damage Ulex europaeus L. (Fabaceae), gorse in the field on mature plants is difficult. Seedlings

may be more affected, given that S. staphylinus at Ulex europaeus was introduced to Hawaii in the high densities in lab experiments can damage and 1800s possibly as a cultivated hedge row plant from Europe, and may also have been used as a graz- kill gorse seedlings (Fowler and Griffin, 1995). ing plant for sheep. It now occupies over 2,000 Populations of this agent may be limited by predators ha on Mauna Kea (Hawaii) and over 2,000 ha in (Markin and Conant, in these proceedings). the Olinda area on Maui. The first biological con- trol effort against gorse was launched by HDOA Tetranychus lintearius Dufour in 1927 and more recently has been a coopera- (Tetranychidae), gorse spider mite tive effort among HDOA, USFS and Landcare Re- First released in 1995 (Culliney and Nagamine, search of New Zealand. In total, eight agents 2000), Tetranychus lintearius dispersed rapidly have been released, but only four became estab- throughout the gorse infestation on Hawaii Island, lished (Markin and Conant, in these proceedings). and was subsequently released on Maui. T. lintearius populations increased tremendously during summer Agonopterix ulicetella (Stainton) (Oecophoridae) seasons, and resulted in considerable damage to Agonopterix ulicetella was first released in gorse foliage in disjunct patches. T. lintearius 1988 and became established on both Hawaii feeding induces mottled chlorosis with stunting, and Maui (Markin et al., 1995). This species and flowering may be aborted on damaged shoots is univoltine and overwinters as adults. In the in spring. Some shoots may even die from heavy spring larvae feed on terminal shoots, sometimes feeding (Hill et al., 1991). However, after the year causing considerable damage, which can reduce 2000, annual T. lintearius population explosions

XIII International Symposium on Biological Control of Weeds - 2011 Session 5 Prospects for Weed Biological Control in Pacific Islands 235 ceased, presumably caused by an accumulation of the island of Hawaii. Proceedings of the Hawaiian predacious mite populations. Entomological Society 32, 123–130. Chun, M.E. (2002) Biological control of ivy gourd, Pempelia genistella (Duponchel) Coccinia grandis (Cucurbitaceae), in Hawaii. In (Pyralidae), gorse hard shoot moth Proceedings of Workshop on Biological Control Releases of Pempelia genistella on Hawaii were of Native Ecosystems in Hawaii (eds Smith, C.W., Denslow, J. and Hight, S.), pp. 8–12. Pacific conducted by USFS 1996-1997, but after initial Cooperative Studies Unit, University of Hawaii at establishment all populations apparently were Manoa, Department of Botany, Technical Report eradicated as a result of a 2001-2002 control effort 129, Honolulu, Hawaii USA. 122p. using herbicide and burning (Markin et al., 2002; Conant, P. (2002) Classical biological control of Markin and Conant, in these proceedings). In its Clidemia hirta (Melastomataceae) in Hawaii using native habitat in Europe, larvae feed during fall multiple strategies. In Proceedings of Workshop and winter months, followed by pupation and adult on Biological Control of Native Ecosystems in emergence in early summer. Larvae often feed Hawaii (eds Smith, C.W., Denslow, J. and Hight, gregariously within webbing on older spines and S.), pp. 13–20. Pacific Cooperative Studies Unit, may girdle the terminal shoots (Markin et al., 1996). University of Hawaii at Manoa, Department of Botany, Technical Report 129, Honolulu, Hawaii Uromyces pisi (DC.) Otth f. sp. europaei Wil- USA. 122p. son and Henderson (Pucciniaceae) Conant, P. (2009) Clidemia hirta (L.) D. Don This rust fungus was released on Hawaii (Melastomataceae) In Biological Control Island in 2000, and was seen once soon after, of Tropical Weeds Using (eds but never recorded again during occasional Muniappan R., Reddy, G.V.P. and Raman, A.). searches over several years (Markin et al., 2002; Cambridge University Press, Cambridge, UK. Markin and Conant, in these proceedings). 508p. Conant, P., Medeiros, A.C. and Loope, L.L. (1997) Acknowledgements A multiagency containment program for miconia (Miconia calvescens) an invasive tree in Hawaiian We thank the following people for comments, rain forests. In Assessment and Management of suggestions and information provided: Mohsen Plant Invasions (eds Luken, J.O. and Thieret, J.W.), Ramadan, Mann Ko, Galen Kawakami, Glenn pp. 249–254. Springer, New York, USA. 324p. Shishido, Hugh Gourlay, Jean-Yves Meyer, Eloise Culliney, T.W. and Nagamine, W.T. (2000) Killgore, Steven Bergfeld, Forest Starr, David Foote Introductions for biological control in and Nicolai Barca. Rachel Winston helped greatly Hawaii,1987–1996. Proceedings of the Hawaiian with review of the manuscript and data in Appendix Entomological Society 34, 121–133. 1. Culliney, T.W., Nagamine, W.T. and Teramoto, K. (2003) Introductions for biological control in References Hawaii 1997–2001. Proceedings of the Hawaiian Entomological Society 36, 145–153. Brenner, G. (2000) Analysis of miconia data. Fowler, S.V. and Griffin, D. (1995) The effects of Hawaii Department of Agriculture, Hilo, Hawaii, multi-species herbivory on shoot growth in USA. Available at: http://www.statpros.com/ gorse Ulex europaeus. Proceedings of the VIII Publications/Miconia_Report.pdf (Accessed 19 International Symposium on Biological Control October 2011). of Weeds (ed Delfosse, E.S.), pp 579–584. DSIR/ Campbell, C.L., Markin, G.P. and Johnson, M.W. CSIRO, Melbourne, . (1995) Fate of Cyanotricha necyri (: Hauff, R. (2006) Controlling banana poka on Maui. Notodontidae) and Pyrausta perelegans Na Leo O Ka Aina. Newsletter of the Hawaii (Lepidoptera: Crambidae) released for biological Department of Land and Natural Resources control of banana poka (Passiflora mollissima) on Division of Forestry and Wildlife 2(1), p. 9.

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Hatcher, P.E., Hinz, H.L. and Rector, B.G.), pp. Proceedings of the Hawaiian Entomological 594-599. CABI Publishing, Wallingford, UK. Society 5, 299-304. Muniappan, R., Reddy, G.V.P. and Raman, A. (2009) Trujillo, E.E. (2005) History and success of plant Coccinia grandis (L.) Voight (Cucurbitaceae). pathogens for biological control of introduced In Biological control of tropical weeds using weeds in Hawaii. Biological Control 33, 113-122. arthropods (eds Muniappan, R., Reddy, G.V.P. and Trujillo, E.E., Kadooka, C., Tanimoto, V., Bergfeld, S., Raman, A.), pp. 175-182. Cambridge University Shishido, G. and Kawakami, G. (2001) Effective Press Cambridge, UK. biomass reduction of the invasive weed species Ramadan, M., Markin, G.P. and Johnson, M.W. banana poka by Septoria leaf spot. Plant Disease (2008) Field biology of Pyrausta perelegans 85, 357-361. (Lepidoptera: Crambidae) a flower bud feeder on Wagner, W.L., Herbst, D.R. and Sohmer, S.H. Passiflora mollissima an invasive vine in Hawaiian (1990) Manual of the Flowering Plants of Hawaii. forests. Tropical Lepidoptera Research 18, 71-79. University of Hawaii Press, Honolulu, Hawaii Reimer, N.J. and Beardsley, J.W. (1988) Effectiveness USA. 988p. of Liothrips urichi Karny (Thysanoptera:Williams, P.A. and Hayes, L. (2007) Emerging weed Phlaeothripidae) introduced for biological control issues for the West Coast Regional Council and of Clidemia hirta in Hawaii, U.S.A. Environmental their prospects for biocontrol. Landcare Research Entomology 18, 1141-1146. Contract Report: LC0607/109. Landcare Reimer, N.J. and Beardsley, J.W. (1986) Some Research, Auckland, New Zealand. 41p. notes on parasitization of Blepharomastix Whiteaker, L.D. and Gardner, D.E. (1985) The ebulealis (Guenee) (Lepidoptera: Pyralidae) distribution of Myrica faya Ait. in the State of in Oahu forests. Proceedings of the Hawaiian Hawaii. Cooperative National Park Resources Entomological Society 27, 91-93. Study Unit, Technical Report 55. Department of Smith, C.W. (2002) Forest pest biological control Botany, University of Hawaii, Honolulu, Hawaii, program in Hawaii. In Proceedings of Workshop USA. 31p. on Biological Control of Native Ecosystems in Whiteaker, L.D. and Gardner, D.E. (1992) Fire tree Hawaii (eds. Smith, C.W., Denslow, J. and Hight, (Myrica faya) distribution in Hawaii. In Alien S.), pp. 91-102. Pacific Cooperative Studies Unit, Plant Invasions in Native Ecosystems of Hawaii: University of Hawaii at Manoa, Department of Management and Research (eds Stone, C.P., Smith, Botany, Technical Report 129, Honolulu, Hawaii C.W. and Tunison, J.T.), pp. 225-240. Cooperative USA. 122p. National Park Resources Studies Unit, University Swezey, O.H. (1924) Records of introduction of of Hawaii, Honolulu, Hawaii USA. 903p. beneficial insects into the Hawaiian Islands.

XIII International Symposium on Biological Control of Weeds - 2011 238 Session 5 Prospects for Weed Biological Control in Pacific Islands 2 Impact Partial Partial established Not Ineffective Ineffective Ineffective Ineffective Partial Partial established Not established Not established Not established Not Partial Substantial Substantial Partial Partial Ineffective Uncertain Ineffective established Not established Not established Not Busck (Froggart) bazochii gundlachianus (Bates) gundlachianus bazochii , 2 spp. Agent Species Agent Octotoma scabripennis Guérin-Méneville Busck lantanella Cremastobombycia sp. Hepialus Strymon echion (L.) Tmolus Lantanophagapusillidactyla (Walker) Aldrich xanthochaeta Eutreta Stål scrupulosa Teleonemia lantana Crocidosema Octotoma gundlachi Suffrain Pseudopyrausta- santatalis (Barnes & McDun nough) Octotoma Baly championi tigris Guenée Diastema (Guenée) sunia Neogalia Salbia haemorrhoidalis Guenée Walker laceratalis Hypena Plagiohammus spinipennis (Thomson) Uroplata girardi Pic Drake Leptobyrsa decora Septoria sp. Ophiomyia lantanae Apion Parevander xanthomelas (Guérin-Méneville) Bates championi Aerenicopsis Agent Family Agent Chrysomelidae Gracillariidae Hepialidae Lycaenidae Lycaenidae Pterophoridae Tephritidae Tingidae Tortricidae Chrysomelidae Crambidae Chrysomelidae Noctuidae Noctuidae Crambidae Noctuidae Cerambycidae Chrysomelidae Tingidae Mycosphaerellaceae Agromyzidae Brentidae Cerambycidae Cerambycidae ; 1953 1 First ReleaseFirst 1902 1902 1902 1902 1902 1902 1902; 1952; 1953; 1954; 1954 1902 1953 1953; 1965 1954 1954; 1962 1955 1956 1957; 1965 1960 1961 1970 1997 1902 1902 1902 1902; 1955 1902 Agent Origin Agent Mexico Mexico Mexico Mexico Mexico Mexico Belize;- Cuba; Flori Mexico; Brazil da, USA; Trinidad; Mexico Cuba Mexico Honduras Mexico Panama; California USA; Cuba Florida, Philippines Kenya/; Mexico Brazil via Australia Colombia/Peru Ecuador Mexico Mexico Mexico Mexico Mexico Target Weed Target Lantana Lantana camara L. (Verbenaceae) Appendix 1. Agents introduced to Hawaii for biological control of weeds 1902-2010, compiled from Julien and Griffiths Appendix 1. Agriculture. and unpublished records of the Hawaii Department (1998), recent publications cited in this review,

XIII International Symposium on Biological Control of Weeds - 2011 Session 5 Prospects for Weed Biological Control in Pacific Islands 239 3 2 Impact Partial established Not established Not Partial Uncertain Ineffective Ineffective Partial established Not established Not Substantial established Not Substantial established Not established Not established Not Substantial Partial (Kirby) Benjamin Stone Walker (Grote) Agent Species Agent Exapion ulicis (Forster) ulicis Exapion sp.Apion Stenopterapion scutellare (Stainton) ulicetella Agonopterix Sericothrips staphylinus Haliday Athesapeuta cyperi Marshall Athesapeuta Bactra venosana (Zeller) lintearius Dufour Tetranychus (Duponchel) genistella Pempelia Wilson pisi europaei (DC.)Uromyces sp. Otth f. Henderson and utilis Procecidochares connexionis Xanthaciura Dactylopius (Cockerell) opuntiae dentata Melitara prodenialis Melitara LeConte armatum Moneilema Cactoblastis cactorum (Berg) Thomson funestus Lagocheirus Agent Family Agent Brentidae Brentidae Brentidae Oecophoridae Thripidae Curculionidae Curculionidae Tortricidae Tetranychidae Pyralidae Pucciniaceae Tephritidae Tephritidae Dactylopiidae Pyralidae Pyralidae Cerambycidae Pyralidae Cerambycidae ; 1955 1 ; 1949 1 First ReleaseFirst 1958 1961; 1989; 1990; 1991 1988; 1991 1991; 1992; 1992 1925 1925 1927 1995 1996 2000 1945 1955 1949 1949 1949 1950 1950 1951 Agent Origin Agent Portugal France Spain; Portugal; Portugal England; France Portugal; England; Philippines Philippines via New England England; Zealand; France via Spain Portugal, England, Zealand viaNew Oregon, USA Portugal England Mexico Mexico via Australia Mexico USA Texas, USA Texas, USA Texas, via Australia Argentina via Australia Mexico

Target Weed Target Purple nutsedge L. rotundus Cyperus (Cyperaceae) Gorse L. europaeus Ulex (Fabaceae) pamakani Maui adenophora Ageratina & H. Rob. King (Spreng.) (Asteraceae) Prickly pear Opuntia spp. (Cactaceae)

XIII International Symposium on Biological Control of Weeds - 2011 240 Session 5 Prospects for Weed Biological Control in Pacific Islands 2 Impact Ineffective Ineffective established Not Ineffective Ineffective Uncertain Ineffective Ineffective Substantial established Not established Not Partial Ineffective Partial Ineffective established Not Partial Ineffective - ) (Wals Hodges Walsingham Clarke Meyrick (Warner) (Snow) Napp atronotatus (Pic) Agent Species Agent Episimus unguiculus Lithraeus Crasimorpha infuscata Strepsicrates smithiana schinella nr. (= C . sp. coruscans Caloptilia Septoria Gardner hodgesii D.E. ingham) Dichomeris aenigmatica (Clarke) Acinia picturata Perapion antiquum (Gyllenhal) antiquum Perapion Perapion neofallax (Kirby) violaceum Perapion Liothrips urichi Karny Ategumia matutinalis Guenée clidemiae f. gloeosporioides sp. C olletotrichum Trujillo poseidon Lius Carposina bullata Meyrick trithalama Mompha Hübner acclinalis Antiblemma Agent Family Agent Tortricidae Bruchidae Gelechiidae Tortricidae Gracillariidae Mycosphaerellaceae Gelechiidae Tephritidae Brentidae Brentidae Brentidae Phlaeothripidae Crambidae Glomerellaceae Buprestidae Carposinidae Coleophoridae Noctuidae First ReleaseFirst 1954 1960 1961 1956 1991 1997 1957 1959 1957 1962 1962 1953 1970 1986 1988 1995 1995 1995 Agent Origin Agent Brazil Brazil Brazil USA Florida/Georgia, Portugal Azores/Madeira, USA Carolina, North Mexico Guatemala South AfricaSouth Morocco Portugal Trinidad via Fiji Trinidad Puerto Rico Trinidad; Panama Trinidad Tobago Tobago Tobago

) faya Emex Emex Raddi Firetree Firetree Sourbush Sourbush (Myricaceae) Koster’s curse Koster’s (Aiton) Wilbur Wilbur (Aiton) (Polygonaceae) (Anacardiaceae) Christmas berry (Melastomataceae) Pluchea carolinensis Schinus terebinthifolius Morella (= Myrica Steinheil australis Emex (L.) Campd. spinosa Emex Don (L.) D. hirta Clidemia (Jacq.) G. Don (Asteraceae) G. Don (Jacq.) Target Weed Target

XIII International Symposium on Biological Control of Weeds - 2011 Session 5 Prospects for Weed Biological Control in Pacific Islands 241 Impact Ineffective Substantial Substantial Ineffective Ineffective Partial Partial established Not Partial Ineffective established Not Substantial Substantial Partial Partial Substantial Not established Not established Not (Hampson) (Hampson) Toll Meyrick (Jacquelin du Val) du (Jacquelin adipalis (Lederer) fatualis (Lederer) Agent Species Agent Loew obscuriventris Tetraeuaresta lareynii Microlarinus Microlarinus lypriformis (Wollaston) Ategumia Rhynchopalpus brunellus Schreckensteinia festaliella Hübner (Harris) Pennisetia marginata Clarke zimmermani Croesia Priophorus morio (Lepeletier) (Fabricius) gibbosa Chlamisus Ategumia (Keiffer) giardi Zeuxidiplosis (Suffrain) quadrigemina Chrysolina & Heppner Yano beneficus Oidaematophorus Steyskal alani Procecidochares & Evans Barreto ageratinae Entyloma Coleophora klimeschiella Coleophora parthenica Agent Family Agent Tephritidae Curculionidae Curculionidae Crambidae Noctuidae Schreckensteiniidae Sesiidae Tortricidae Tenthridinidae Chrysomelidae Crambidae Cecidomyiidae Chrysomelidae Pterophoridae Tephritidae Entylomataceae Coleophoridae Coleophoridae First ReleaseFirst 1961 1962 1963 1965 1965 1963 1963 1964 1966 1969 1958 1965 1965 1973 1974 1975 1980 1980 - Agent Origin Agent Trinidad via Fiji Trinidad via USA Calif./Arizona, Italy via USA Calif./Arizona, Italy California, USA Malaysia Malaysia USA Oregon, Mexico USA Oregon/California, USA Missouri, Philippines France via California, USA France Zealand via Australia/New France via Australia via via Australia France California, USA Mexico Mexico Jamaica Pakistan via California, USA Pakistan via Califor Egypt/Pakistan nia, USA

Lour. Lour. (Rosaceae) (Asteraceae) Puncture vine vine Puncture Klamath weed Klamath Elephant’s foot foot Elephant’s (Hypericaceae) Russian thistle Russian Ageratina riparia riparia Ageratina (Zygophyllaceae) Asian melastome Asian Salsola tragus L. tragus Salsola Florida blackberryFlorida (Melastomataceae) Rubus argutus Link Kunth (Asteraceae) Kunth Tribulus cistoidesTribulus L. Elephantopus mollis Tribulus terrestrisTribulus L. Hamakua pamakani pamakani Hamakua (Chenopodiaceae) Hypericum perforatum perforatum Hypericum (Regel) King & H. Rob. & H. Rob. King (Regel) Melastoma septemnervium Target Weed Target

XIII International Symposium on Biological Control of Weeds - 2011 242 Session 5 Prospects for Weed Biological Control in Pacific Islands Impact Partial Not established Not Ineffective Partial Substantial established Not Partial - ) necyria- & Rogen (Felder Agent Species Agent Killgore et al. Killgore coniae mi sp. f. (Penz.) gloeosporioides Colletotrichum Scea (= Cyanotricha Hampson perelegans Pyrausta Septoria passiflorae Syd. hofer) Melittia oedipus Oberthür oedipus Melittia Acythopeus burkhartorum Pakaluk and O’Brien Pakaluk Acythopeus and cocciniae O’Brien Agent Family Agent Glomerellaceae Notodontidae Crambidae Mycosphaerellaceae Sesiidae Curculionidae Curculionidae First ReleaseFirst 1997 1988 1991 1996 1996 1999 1999 Agent Origin Agent Brazil Colombia/ Ecuador Colombia/ Venezuela Colombia Kenya Kenya Kenya

Miconia (L.) Voigt (L.) Voigt Ivy gourd gourd Ivy Banana poka (Passifloraceae) (Cucurbitaceae) Coccinia grandis grandis Coccinia (Melastomataceae) Passiflora tarminiana Miconia calvescens DC. calvescens Miconia Coppens & V.E. Barney & V.E. Coppens These early releases failed to establish. Established, but population and/or distribution limited, and evidence on impact lacking. Ageratina spp. Agent apparently was released on both Target Weed Target 1 2 3

XIII International Symposium on Biological Control of Weeds - 2011