FORUM Accumulation of Pest Insects 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 insect 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 genus 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 Trachymela 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 Aprostocetus sp. 19951 200017 Epichrisocharis burwelli 19996 Selitrichodes 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 gall 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