FORUM Accumulation of Pest on Eucalyptus in California: Random

Process or Smoking Gun Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019

1,2 1 3 TIMOTHY D. PAINE, JOCELYN G. MILLAR, AND KENT M. DAANE

J. Econ. Entomol. 103(6): 1943Ð1949 (2010); DOI: 10.1603/EC10214 ABSTRACT Eucalyptus spp., native to Australia, have been introduced into many parts of the world as important timber and ornamental trees. Although the trees have important silvicultural qualities, they also have generated intense dissatisfaction, particularly among groups of individuals in California. The trees have beneÞted from the lack of pests and diseases in their adventive ranges but that has changed over the past four decades. In California, two species of insect herbivores were introduced between the time trees were Þrst introduced to the state in the middle of the 19th century and 1983. Between 1983 and 2008, an additional 16 Australian insect pests of eucalyptus have become established in the state. The modes or routes of introduction have never been established. However, examinations of different temporal and spatial patterns suggest that the introductions were nonrandom processes. It is possible that they occurred because of increased trade or movement of people, but the hypothesis that there were intentional introductions also must be considered. The rapid accumulation of intro- duced herbivores on an ornamental plant system in a single state is a cautionary example of what could happen if a major food or Þber crop were intentionally targeted.

KEY WORDS Eucalyptus, invasive species, intentional introductions, insect herbivores, adventive range

Eucalyptus spp. are native to Australia and New in Arizona, Florida, Hawaii, and into Mexico for similar Guinea (Doughty 2000). They have earned a unique purposes. Limited in distribution by low temperatures, reputation among timber species because of several they grow well under hot, dry summer conditions and desirable characteristics. For example, many species mild winters, often where other ornamental trees do grow rapidly and produce very high-quality wood Þ- not thrive. ber that can be used as timber or as cellulose stock for In addition to their rapid growth, relatively high manufacturing high-quality paper for laser printers drought tolerance, and ability to grow on poor soils, and copy machines. They can be planted at high den- the widespread adoption of Eucalyptus was enhanced sities in plantations and can be harvested at rotations by the lack of insect pests in most of the introduced of Ͻ10 yr, depending on location. The trees will grow ranges. The trees were widely distributed around the on poor soils with no supplemental irrigation, giving world from Australia in the mid-19th century at the them an advantage over other hardwood plantation end of the age of sail and the beginning of the era of species. Consequently, vast plantations of Eucalyptus steam ships. The long voyages made transportation of are found in South America (Brazil, Chile, and Ar- large numbers of live plants impractical. Fortunately, gentina), sub-Saharan Africa (Republic of South Af- the trees propagate easily from seed. One of the con- rica, Zambia, and Malawi), and the Mediterranean sequences of seed propagation is that the foliage-, sap-, basin (Spain, Portugal, Italy, Tunisia, Algeria, and Is- and wood-feeding insects were not moved with the rael) (Doughty 2000). seeds (Doughty 2000). Besides their value as commercial species, Eucalyp- Although the trees have signiÞcant commercial and tus also have been planted in many parts of the tem- esthetic value, they also have received criticism and perate and subtropical world as ornamental trees. labeled as an undesirable or invasive species. Many They were introduced into California initially as a species have exfoliating bark and, although evergreen, timber species, but the growth characteristics were they can shed large amounts of leaf litter and seed unsuitable. However, 90 species in the have capsules that many Þnd messy and a nuisance. If the been widely planted as landscape and windbreak trees shed litter is allowed to accumulate under the trees, it across California (Doughty 2000) and are also grown can function as a critical fuel source for urban wild- Þres. In addition, the leaves have high levels of ter- 1 Department of Entomology, University of California, Riverside, penes, responsible for the characteristic Eucalyptus CA 92521. odor and a source of Eucalyptus oil, that make the 2 Corresponding author, e-mail: [email protected]. 3 Department of Environmental Science, Policy, and Management, foliage highly ßammable. The rapid growth also can University of California, Berkeley, CA 94720. enable some naturalized species to invade native land-

0022-0493/10/1943Ð1949$04.00/0 ᭧ 2010 Entomological Society of America 1944 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 103, no. 6

Table 1. Date of first report of invasive species feeding on Eucalyptus in four adventive regions of the world

Region Species North South Southern

Europe/Mediterranean Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019 Americaa America Africa Phoracantha semipunctata 19841 19409 Ϸ191021 190628 Phoracantha recurva 19951 199210 197021 194928 Gonipterus scutellatus 19941 197911 192622 191629 sloanei 19981 197630 Trachymela tinticollis 198230 Chrysophtharta m-fuscum 20032 Acizzia uncatoidesb 19543 Blastopsylla occidentalis 19831 199723 Ctenerytaina longicauda 19831 Ctenerytaina eucalypti 19911 199812 199922 Ctenerytaina spatulata 19911 200213 199424 Ctenerytaina peregrina 200714 Cryptoneossa triangula 19951 Glycaspis brimblecombei 19981 200715 200325 Eucalyptolyma maideni 20004 Quadrastichodella nova 19575 197716 195716 197716 sp. 19951 200017 Epichrisocharis burwelli 19996 globulus 20087 Leptocybe invasac 20088 200018 Leprosa milga 200619 200319 Ophelinus maskelli 200020 Megastigmus eucalypti 200017 Megastigmus zebrinus 200631 Moona spermatophaga 200026 200819 Thaumastocoris peregrinus 200627 200532

a Unless otherwise noted, all of the initial reports from North America are from California. b The primary host of this psyllid is Acacia sp. but has been reported colonizing Eucalyptus sp. c The only reported occurrence of this forming wasp in North America is from Florida. 1Gill (1998), 2vonEllenreider (2003), 3Ulyshen and Miller (2007), 4Garrison and vonEllenreider (2003), 5Timberlake (1957), 6Schauff and Garrison (2000), 7La Salle et al. (2009), 8Willey and Skelley (2008), 9Mendel (1985), 10BenJamaa et al. (2002), 11Rabasse and Perrin (1979), 12Burckhardt (1998), 13Valente et al. (2004), 14Hodkinson (2007), 15Valente and Hodkinson (2009), 16La Salle (2005), 17Viggiani et al. (2002), 18Mendel et al. (2004), 19Kim and La Salle (2008), 20Protasov et al. (2007), 21Di-lorio (2004), 22Lanfranco and Dungey (2001), 23Santana and Burckhardt (2007), 24Taylor (1997), 25Firmino-Winckler et al. (2009), 26Kim et al. (2005), 27Carpintero and Dellape (2006), 28Drinkwater (1975), 29Tooke (1955), 30Tribe and Cillie (1997), 31Grissell (2006), 32Jacobs and Neser (2005). scapes. The detractorsÕ perception is that that some free, with a single exception. That exception was a eucalypt species are partly responsible for changing seed-galling eulophid wasp Quadrastichodella nova the natural landscape by crowding out or preventing Girault (Timberlake 1957) that could have arrived in the growth of native species. Eucalyptus globulus La- early shipments of seeds. Also reported during this billardie`re, Tasmanian blue gum, is classiÞed as a po- period was the psyllid Acizzia uncatoides (Ferris & tentially harmful invasive species by the California Klyver), which was introduced in 1954 (Ulyshen and Exotic Plant Council, a nongovernmental organization Miller 2007). This psyllid is primarily a pest of Acacia (California Invasive Plant Council 2006). Some writ- spp., but on rare occasions it will feed on some Euca- ers have gone so far as to suggest that introduced lyptus spp. Between 1984 and 2008, an additional 15 herbivores are potential biological control agents of species in four different guilds (two species of cole- Eucalyptus in California (La Salle et al. 2009). In some opteran wood borers, three species of coleopteran cases in countries with large commercial plantations, defoliators, seven species of ßuid feeding psyllids, and the trees can symbolize widespread conversion of the three species of hymenopteran gallers) were discov- landscape from traditional uses and replacement of ered in the state (Table 1). All of these insects are native plant communities. However, it is unclear Australian in origin. whether this situation is any different than the large How did these insect pests arrive in the state in such scale cultivation of any of the major crop or forestry a short time frame? Was this simply a matter of in- species that are crucial to providing adequate food and creased movement of people and goods across the shelter for the worldÕs current and future populations. PaciÞc Ocean in the last two and a half decades or has there been an intentional movement of plant pests? The question troubles landscape designers, arborists, The California Question and nursery producers in the state because it affects The Þrst Eucalyptus trees planted in North America pest management operations and choices about what were planted in southern California around 1850. Be- to plant and what to produce. On a much broader tween 1850 and 1983, a period spanning nearly a cen- level, whereas the introduced Eucalyptus are not vital tury and a half, Eucalyptus were reported to be pest to national security in terms of food, Þber, or timber December 2010 PAINE ET AL.: PEST INSECTS ON EUCALYPTUS IN CALIFORNIA 1945

Table 2. Native geographic range in Australia of invasive pests of Eucalyptus introduced into California between 1983 and 2000 (ABRS 2009)

Australian state

New Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019 Year and species South Western Northern South Queensland Victoria Tasmania Australia Australia Territory Wales 1983Ð1984 B. occidentalis XXXX C. longicauda X P. semipunctata XXXXXXX 1990Ð1991 C. eucalypti XXXX XX C. spatulata XX X X 1994Ð1995 G. scutellatus XX X X X P. recurva XXXXXXX C. triangula XXXX 1998Ð1999 G. brimblecombei XX X T. sloanei X E. burwellia 2000 E. maideni XX X

a Data are unavailable. production, the question also should pique the interest into the state: 1983Ð1984 [Phoracantha semipunctata of those agencies responsible for regulating interna- (F.), Blastopsylla occidentalis (Taylor), and Ctenar- tional movement of goods and people. ytaina longicauda Taylor], 1990Ð1991 (C. eucalypti It is important to recognize at the outset that there and Ctenarytaina spatulata Taylor), 1994Ð1995 (Pho- is no single answer to the question. No group or in- racantha recurva Newman, Gonipterus scutellatus dividual has claimed any responsibility for introducing Gyllenhal, Cryptoneossa triangula Taylor, and Apros- exotic insects from Australia into California. What we tocetus sp.), and 1998Ð1999 [Trachymela sloanei have is a series of patterns that can be examined. There (Blackburn), Glycaspis brimblecombei Moore, and are a number of possible alternative explanations for burwelli Schauff]. An additional spe- each pattern; however, the conclusions drawn from cies detected in 2000, Eucalyptolyma maideni Frog- those patterns are left to the reader to interpret. gatt, immediately after this last period. Other than the exception noted, the detections were localized to a single geographic areaÑalthough eucalypts are Patterns of Introductions planted throughout CaliforniaÑand, for the last three Table 1 lists Eucalyptus-feeding insects that have clusters, in narrow time periods separated by 3-yr been introduced into four different eucalypt growing breaks (Table 1). regions of the world and the year in which each insect If there is a temporal relationship for groups of was reported from that region. The year an insect was species introduced into California, there also may be Þrst reported is used as a proxy for the year of intro- a spatial pattern in their distribution in their native duction, but it must be clearly stated that they are not range. That is, collection of species for simultaneous necessarily the same thing. The time required for group introductions would be facilitated if the species populations to increase to detectable levels, the num- had overlapping geographic ranges in Australia. The ber of trained observers in a region, and the systematic three species detected in 1983Ð1984 are all found in resources available to recognize and identify newly Queensland (Table 2) (ABRS 2009). Similarly, the two introduced species can all inßuence any temporal gap species detected in 1990Ð1991 are both found in New between introduction and detection of an invasive South Wales and South Australia, whereas the four pest on Eucalyptus. species detected in 1994Ð1995 are found in those two There are several local patterns that emerge from states and in Queensland (Table 2). The three species examination of the California introductions. With the detected in 1998Ð1999, as well as the one species exception of the psyllid Ctenarytaina eucalypti detected in 2000, are all found in New South Wales Maskell, all of the introductions were detected Þrst in (Table 2). the southern part of the state. In contrast, C. eucalypti The spatial and temporal patterns of introduction in was Þrst reported in commercial eucalypt foliage pro- California do not seem to be random, particularly duction in Monterey County and may have been as- when taken in perspective of the geographic distri- sociated with movement of commercial cultivars. In butions of the insects in Australia. Although there are addition to the spatial pattern of introduction, there is many potential routes of introduction, there are two a temporal pattern. There were four periods between that would seem to stand out. The Þrst is importation 1983 and 1998 when multiple species were introduced of infested plant material. California has the nationÕs 1946 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 103, no. 6 largest wholesale nursery industry and production is Eucalyptus growing regions. There are eight species centered south of the San Francisco Bay area and in found in other parts of the world but not in California; the southern California counties. However, because of Þve of those species are seed galling and the controversial nature of Eucalyptus in the state, the probably moved in infested seed lots as plantations of Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019 nursery industry has not expanded the range of cul- Eucalyptus were expanded worldwide (e.g., Moona tivars of Eucalyptus used in California in many years. spermophaga Kim & La Salle) (Kim et al. 2005). Seven That is, production and sale is limited to species and of the species found outside California were Þrst de- cultivars already established in the state and not on tected in their adventive ranges since 2000, after the imported material from Australia. Therefore, it is un- period of greatest invasive activity in California had likely that the insects, particularly those that have past. Of the seven species found only in California, Þve immature stages that require live host material, were of them are psyllids that require living plant material introduced through commercial trafÞc of Eucalyptus to support nymphal survival. Again, the pattern is not plants moving from Australia into California. random across feeding guilds. The second is international movement of goods and Wood borers and leaf are robust and cryptic, people through transportation hubs. The ports of Oak- whereas psyllids are soft bodied, have short lives, and land on the San Francisco Bay in northern California require living hosts in their immature stages (Paine and the ports of Los Angeles and Long Beach in south- and Millar 2002, Daane et al. 2005). The diversity of ern California are among the busiest container ship- psyllids in California suggests either movement of ping ports in the country. Similarly, San Francisco plant material or protected movement of adults. If the International and Los Angeles International airports psyllids were intentionally introduced, the level of are major North American points of entry for passen- success suggests either a signiÞcant base of knowledge gers and air cargo. Despite the similarities in centers of insect biology and life history, someone who was of international commerce between the geographic remarkably lucky at smuggling in live insects and plant regions, all but one of the introductions have been in materials for those insects that will not survive without the south. hosts without getting caught, or someone who does Statistics compiled by the California Department of not have to go through the normal customs and agri- Food and Agriculture up to 2006 (R. V. Dowell, per- cultural inspection procedures, such as someone trav- sonal communication) show that 28 insect herbivores eling with a special passport or the equivalent. Alter- native to Australia had established in California and natively, it could possibly be done by someone who is that 16 feed on Eucalyptus. An additional Eucalyptus shipping in a large volume of plant material, such as a specialist was collected in 2008 and identiÞed in 2009 commercial nursery, because only a small percentage (La Salle et al. 2009). From the time that records were of each shipment can probably be checked thor- Þrst kept until the decade of the 1980s, 11 Australian oughly. species had invaded the state, including the Host range patterns of the introduced Eucalyptus two previously mentioned (Q. nova and A. uncatoides) pests also suggest skewed invasion events. Although that used Eucalyptus as a host. That pattern changed there are some 90 species of Eucalyptus present in dramatically in the next three decades. In the 1980s, California, the more serious wood borer, leaf , three of the four introduced insects from Australia fed and psyllid pests species found in the state colonize, on Eucalyptus, nine of 11 introduced from that con- among other species, the Tasmanian blue gum and tinent in the 1990s used Eucalyptus as a host, and both river red gum, Eucalyptus camaldulensis Dehnh. Both of the Australian insects detected in the Þrst half of the of these species are abundant in the state and have 2000s were speciÞc to Eucalyptus. There are many suffered signiÞcant levels of mortality as a result of agricultural and commercial products derived from insect activity (Paine and Millar 2002, Daane et al. many different plant genera produced in Australia and 2005). They are also the only two Eucalyptus species traded in California, and yet 14 of the 17 Australian listed on the invasive plant inventory by the California insects that established in the state after 1980 were Invasive Plant Council (California Invasive Plant speciÞc to a single plant genus. The pattern seems to Council 2006). be nonrandom. Another pattern that accompanied the introduction Eucalyptus are grown in many parts of the world and of Eucalyptus pests into California was the lack of their these regions have also accumulated insects from Aus- associated natural enemies. The concept of propagule tralia (Table 1). The international pattern of intro- pressure in invasion ecology suggests a greater chance ductions provides a contrast to the pattern in Califor- for establishment success when the invasive species nia. There are 25 species of herbivores that have been are released in higher numbers into the non-native introduced from Australia to various parts of the world environment (Lockwood et al. 2005, Von Holle and (Table 1). In total, 17 pest species are found in Cali- Simberloff 2005). It is likely, therefore, that any acci- fornia; 10 of those species are shared with at least one dental establishment of the eucalypt insect pest spe- other geographic region and seven species are found cies in California involved movement of large amounts only in California. However, six of the shared species of infested eucalypt foliage or timber. were Þrst recorded from California. The remaining Many eucalypt herbivores in at least some of the four species (two wood borers and two leaf beetles) invasion events would imply the associated importa- were very early introductions into South Africa, and tion of their specialized natural enemies. For example, three species are now distributed throughout all of the in the Isle of Man (British Crown Dependency), the December 2010 PAINE ET AL.: PEST INSECTS ON EUCALYPTUS IN CALIFORNIA 1947 invasion by C. eucalypti was accompanied by its para- be eradicated. That is, rather than controlling the sitoid Psyllaephagus pilosus Noyes (Bennett 2005). In herbivores, a vocal minority has suggested that inva- their native Australian-Tasmanian ranges, the long- sive insect species that attack Eucalyptus should be horn beetles (Phoracantha spp.), psyllids (Ctenar- encouraged as biological control agents. Although Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019 ytaina spp., G. brimblecombei), and a weevil (G. scute- many members of audiences at our public forums on llatus) are attacked by a suite of specialist natural Eucalyptus pest management presentations would ap- enemies (Riek 1962, Huber and Prinsloo 1990, Austin proach researchers wanting to know how to rear and et al. 1994). Nevertheless, surveys during the initial redistribute natural enemies to eliminate pest species, years after these pests arrived in California found none other individuals at the same events sought informa- of their associate natural enemies had arrived in Cal- tion on how to rear and redistribute pests. ifornia during the initial invasion event, leading to the It must be emphasized that these sentiments are not relatively successful implementation of classical bio- ubiquitous but instead are representative of the per- logical control for these pest species (Hanks et al. ceptions of a relatively small but vocal community 2000; Paine et al. 2001, 2004; Paine and Millar 2002; opposed to non-native plants in California. Whereas Luhring et al. 2004; Daane et al. 2005). The one ex- the patterns are powerful and repetitive, without a ception was the recent importation of E. maideni, from substantiated or documented claim of responsibility which a specialized Psyllaephagus species were recov- they are no more than strongly suggestive. In that ered in California some 4 yr after the pest was Þrst light, the hypothesis that there have been intentional detected (T.D.P. and K.M.D., unpublished data). The introductions of Eucalyptus pests into California may lack of associated natural enemies for so many of the be nothing more than an accumulation of coinci- eucalypt pests in California might imply a prerelease dences and a modern version of a Just So Story selection for only herbivores. (Kipling 1902). Alternatively, we suggest that the ac- Invasion events for Eucalyptus-feeding herbivores cumulation of circumstantial patterns may point to in South America, Europe, and South Africa would not nonrandom, intentional processes with important im- seem to support the notion that specialized natural plications. Although Eucalyptus are not vital to na- enemies would necessarily also accompany the initial tional food or health security, the possibility that these invasion of eucalypt pests (Lanfranco and Dungey introductions, with a cumulative cost of hundreds of 2001, Chauzat et al. 2002, De Marzo 2007). The ap- millions of dollars in direct damage, control measures, pearance of several pest species in those areas without and removal and/or replacement of tens of thousands natural enemies would cast doubt on the hypothesis of killed trees, may be the result of carefully planned that the lack of natural enemies across a broad range and executed introductions of known pests should be of herbivores in California may have been intentional cause for major concern. In particular, the effective- introductions. However, many of these pests appeared ness of these introductions at causing widespread Þrst in California and then more broadly in other parts damage to valuable plant species highlights the ease of the world. If the source of the invasion in those with which a knowledgeable and dedicated group of other sites was from the adventive range without nat- individuals could intentionally introduce ecologically ural enemies of the pests (e.g., California) rather than or economically destructive pests and diseases, par- from the native range, then the herbivores would ticularly those of major food crops. escape control. The introduction of exotic organisms into new parts of the world predates human origins, with new colo- nizers arriving by wind or water. With the rise of Summary human civilizations, numerous introductions (e.g., The accumulation of herbivores on Eucalyptus in crop species of plants and domesticated ) have California has attracted concern and interest among been intentional. However, many more have been the arborist, nursery, and regulatory communities. Six- accidental. Accidental introductions of crop or forest teen species arrived in a period of just Ͼ20 yr after pests such as Asian citrus psyllid, Diaphorina citri Ku- Ϸ100 yr without any introductions at all. There are wayama; Asian longhorned borer, Anoplophora gla- temporal and spatial patterns of introduction that bripennis (Motschulsky); emerald ash borer, Agrilus could be coincidence and nothing but the summation planipennis Fairmaire; or soybean aphid, Aphis gly- of random economic or population-driven processes. cines Matsumura, into North America in recent years In fact, introductions of insects affecting urban and have illustrated the environmental damage and accu- forest trees are mounting around the world (Niemela mulated direct costs that can accrue in trying to erad- and Mattson 1996). icate, mitigate, or manage these invasive pest species Alternatively, but with no deÞnitive proof, we sug- and their associated pathogens. Intentional introduc- gest that the multiple patterns may be nonrandom; tions of insect herbivores onto crop plants, or organ- instead, they suggest the possibility of intentional in- isms pathogenic to plants or domestic animals, repre- troductions. Eucalyptus have been the focus of intense sent an insidious threat that could severely damage the dislike in many parts of California. It has not been national agricultural economy, endanger a safe and uncommon to be confronted with challenges to bio- abundant food supply, threaten water quality or quan- logical control research programs (e.g., P. semipunc- tity, increase the risk of wild Þres, or degrade envi- tata, Hanks et al. 1995 or G. scutellatus, Hanks et al. ronmental quality across massive areas. In the absence 2000) for wasting resources to save trees that should of a claim of responsibility, examination of the patterns 1948 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 103, no. 6 of species accumulation can provide crucial clues as to Garrison, R. W., and N. vonEllenreider. 2003. Spotted gum sources of introductions, and whether they were ac- lerp psyllid (Eucalyptolyma maideni). California Depart- cidental or, potentially, intentional. ment of Agriculture Pest Sheets. CDFA, Sacramento, CA. Gill, R. J. 1998. Recently introduced pests of Eucalyptus. Calif. Plant Pest Dis. Rep. 17: 21Ð24. Downloaded from https://academic.oup.com/jee/article-abstract/103/6/1943/2199474 by Cal Polytechnic State Univ San Luis Obispo user on 09 June 2019 Acknowledgments Grissell, E. E. 2006. A new species of Megastigmus Dalman (Hymenoptera: Torymidae), galling seed capsules of Eu- The research was supported, in part, by grants from the calyptus camaldulensis Denhardt (Myrtacae) in South CSREES/UC Invasive and Exotic Pest and Disease Program Africa and Australia. Afr. Entomol. 14: 87Ð94. (to T.D.P. and J.G.M.). Hanks, L. M., J. R. Gould, T. D. Paine, J. G. Millar, and Q. Wang. 1995. Biology and host relations of Avetianella longoi. (Hymenoptera: Encyrtidae), an egg of References Cited the eucalyptus longhorned borer (Coleoptera: Ceram- bycidae). Ann. Entomol. Soc. Am. 88: 666Ð671. Austin, A. D., D.L.J. Quicke, and P. M. Marsh. 1994. The Hanks, L. M., J. G. Millar, T. D. Paine, and C. D. Campbell. hymenopterous of Eucalyptus longhorn bee- 2000. Classical biological control of the Australian weevil tles, Phoracantha spp. (Coleoptera, Cerambycidae) in Gonipterus scutellatus Gyll. (Coleoptera: Curculionidae) Australia. Bull. Entomol. Res. 84: 145Ð174. in California. Environ. 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