Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Post-establishment assessment of host plant specificity of Arytainilla spartiophila (Hemiptera: Psyllidae), an adventive biological control agent of Scotch broom,Cytisus scoparius

Scotch broom, Cytisus scoparius (Fabaceae), is a shrub native to Europe that is invasive in the USA, New Zealand and Australia. The psyllid Arytainilla spartiophila has been purposely introduced to Australia and New Zealand as a biological control agent of C. scoparius, but is an accidental introduction to California. Lupines (Lupinus spp.) are the closest native taxon to Cytisus in North America, and are therefore considered to be at the highest risk for non-target damage. However, because no lupines are native to Australia or New Zealand, only one imported forage species was evaluated during prior host specificity testing. We conducted a laboratory nymphal transfer experiment, a field choice experiment and a field survey to assess risk to three lupine species (Lupinus albifrons, Lupinus bicolor and Lupinus formosus). In the laboratory, 20% of third-instar nymphs were able to develop to adulthood on L. formosus but not on the other lupine species, while 40% completed development on C. scoparius. In the field experiment, potted lupine and C. scoparius plants were placed beside large infested C. scoparius plants; oviposition occurred on all the potted C. scoparius plants, but on none of the lupines. In the field survey, no A. spartiophila eggs or nymphs were found on naturally occurring lupines growing adjacent to infested C. scoparius. The results indicate that A. spartiophila is not likely to damage or reproduce on lupines in the field. This study provides an example of how field studies can help clarify the host specificity of biological control agents.     

Can enemy release explain the invasion success of the diploid <Emphasis Type="Italic">Leucanthemum vulgare</Emphasis> in North America?

Enemy release is a commonly accepted mechanism to explain plant invasions. Both the diploid Leucanthemum vulgare and the morphologically very similar tetraploid Leucanthemum ircutianum have been introduced into North America. To verify which species is more prevalent in North America we sampled 98 Leucanthemum populations and determined their ploidy level. Although polyploidy has repeatedly been proposed to be associated with increased invasiveness in plants, only two of the populations surveyed in North America were the tetraploid L. ircutianum. We tested the enemy release hypothesis by first comparing 20 populations of L. vulgare and 27 populations of L. ircutianum in their native range in Europe, and then comparing the European L. vulgare populations with 31 L. vulgare populations sampled in North America. Characteristics of the site and associated vegetation, plant performance and invertebrate herbivory were recorded. In Europe, plant height and density of the two species were similar but L. vulgare produced more flower heads than L. ircutianum. Leucanthemum vulgare in North America was 17 % taller, produced twice as many flower heads and grew much denser compared to L. vulgare in Europe. Attack rates by root- and leaf-feeding herbivores on L. vulgare in Europe (34 and 75 %) was comparable to that on L. ircutianum (26 and 71 %) but higher than that on L. vulgare in North America (10 and 3 %). However, herbivore load and leaf damage were low in Europe. Cover and height of the co-occurring vegetation was higher in L. vulgare populations in the native than in the introduced range, suggesting that a shift in plant competition may more easily explain the invasion success of L. vulgare than escape from herbivory.     

Microsatellite isolation from the gall midge Spurgia capitigena (Diptera: Cecidomyiidae), a biological control agent of leafy spurge

Spurgia capitigena is a gall midge that has been released as a biological control agent of leafy spurge (Euphorbia esula), a noxious rangeland weed in North America. We isolated 15 microsatellite loci from S. capitigena with from two to 27 alleles at each locus. These markers will allow us to examine the structure and levels of neutral genetic variation in native and introduced populations.     

Is there rapid evolutionary response in introduced populations of tansy ragwort, <Emphasis Type="Italic">Jacobaea vulgaris</Emphasis>, when exposed to biological control?

Differences in the herbivore community between a plant’s native (specialists and generalists) and introduced range (almost exclusively generalists) may lead to the evolution of reduced allocation to defences against specialist herbivores in the introduced range, allowing for increased allocation to competitive ability and to defences against generalist herbivores. Following this logic, the introduction of biological control agents should reverse this evolutionary shift and select for plants with life-history traits that are more similar to those of plants in the native range than those of plants in the introduced range that have not been exposed to biological control. In a common garden experiment, we compared performance and resistance traits of tansy ragwort, Jacobaea vulgaris, among populations from the introduced range (New Zealand and North America) that have either been exposed to or grown free from the biological control agent Longitarsus jacobaeae. For comparison, we included populations from the native European range. We found lower levels of generalist-deterrent pyrrolizidine alkaloids (PAs) and of soluble phenolics in New Zealand populations with than in populations without exposure to L. jacobaeae, while the opposite pattern was detected among North American populations. Contrary to expectation, populations with exposure to L. jacobaeae revealed more feeding damage by L. jacobaeae than populations without exposure. Introduced populations had higher levels of PAs and reproductive output than native J. vulgaris populations. Jacobaea vulgaris was introduced in different parts of the world some 100–130 years ago, while L. jacobaeae was introduced only some 20–40 years ago. Hence, the larger differences observed between native and introduced populations, as compared to introduced populations with and without biological control history, may result from different time scales available for selection to act.     

Biology and host specificity of Aulacobaris fallax (Coleoptera: Curculionidae), a potential biological control agent for dyer’s woad,Isatis tinctoria (Brassicaceae) in North America

Dyer’s woad, Isatis tinctoria, a plant of Eurasian origin is a problematic weed in western North America against which a classical biological weed control programme was initiated in 2004. Three European insect species were selected as candidate agents to control this invasive species, including the root‐mining weevil Aulacobaris fallax. To determine its suitability as an agent, the biology and host specificity of A. fallax were studied in outdoor plots and in the field between 2004 and 2006 in its native European range. Aulacobaris fallax is a univoltine species that lays its eggs from March to August into leaf stalks and roots of dyer’s woad. Larvae mine and pupate in the roots and adults emerge from August to October. Up to 62% of the dyer’s woad plants at the field sites investigated were attacked by this weevil. In no‐choice host‐specificity tests, A. fallax attacked 16 out of 39 species and varieties within the Family Brassicaceae. Twelve of these are native to North America. In subsequent multiple‐choice tests, seven species, all native to North America, suffered a similar level of attack as dyer’s woad, while none of the European species were attacked. Our results demonstrate the importance of including test plant species that have not co‐evolved with the respective candidate agent. In sum, we conclude that the risk of non‐target effects is too high for A. fallax to be considered as a biological control agent for dyer’s woad in the United States.     

Population genetics and symbiont assemblages support opposing invasion scenarios for the red turpentine beetle (Dendroctonus valens)

Exotic forest insects and their symbionts pose an increasing threat to forest health. This is apparently true for the red turpentine beetle (Dendroctonus valens), which was unintentionally introduced to China, where the beetle has killed millions of healthy native pine trees. Previous population genetics studies that used cytochrome oxidase I as a marker concluded that the source of D. valens in China was western North America. In contrast, surveys of fungi associated with D. valens demonstrated that more fungal species are shared between China and eastern North America than between China and western North America, suggesting that the source population of D. valens could be eastern North America. In this study, we used microsatellite markers to determine population structure of D. valens in North America as well as the source population of the beetle in China. The analyses revealed that four genetically distinct populations (herein named the West, Central, Northeast and Mexico) represent the native range of D. valens. Clustering analyses and a simulation‐based approximate Bayesian computation (ABC) approach supported the hypothesis that western North America is the source of the invasive D. valens population. This study provides a demonstration of non‐congruence between patterns inferred by studies on population genetics and symbiont assemblages in an invasive bark beetle.     

Evaluation of the rust fungus Puccinia rapipes for biological control of Lycium ferocissimum (African boxthorn) in Australia: Life cycle,taxonomy and pathogenicity

Fungal plant pathogens are increasingly recognised as being among the most effective and safe agents in classical weed biological control programs worldwide. Suitability of the rust fungus P. rapipes as a classical biological control agent for Lycium ferocissimum (African boxthorn) in Australia was assessed using a streamlined agent selection framework. Studies with P. rapipes were undertaken to elucidate its life cycle, confirm its taxonomic placement and determine its pathogenicity to L. ferocissimum and seven closely-related Solanaceae species that occur in Australia. Field surveys in the native range of South Africa, experiments in a containment facility in Australia and DNA sequencing confirmed that P. rapipes is macrocyclic and autoecious, producing all five spore stages on L. ferocissimum. The stages not previously encountered, spermogonia and aecia, are described. Sequencing also confirmed that P. rapipes is sister to Puccinia afra, in the ‘Old World Lineage’ of Puccinia species on Lycieae. Two purified isolates of the fungus, representing the Eastern and Western Cape distributions of P. rapipes in South Africa, were cultured in the containment facility for use in pathogenicity testing. L. ferocissimum and all of the Lycium species of Eurasian origin tested ‒ Lycium barbarum (goji berry), Lycium chinense (goji berry ‘chinense’) and Lycium ruthenicum (black goji berry) – were susceptible to both isolates of P. rapipes. The Australian native L. australe and three more distantly related species in different genera tested were resistant to both isolates. The isolate from the Western Cape was significantly more pathogenic on L. ferocissimum from Australia, than the Eastern Cape isolate. Our results indicate that P. rapipes may be sufficiently host specific to pursue as a biological control agent in an Australian context, should regulators be willing to accept damage to the Eurasian goji berries being grown, albeit to a limited extent, in Australia.     

Biology of Platycephala planifrons (Diptera: Chloropidae) and its potential effectiveness as biological control agent for invasive Phragmites australis in North America

Phragmites australis is a cosmopolitan clonal grass valued for its support of diversity-rich communities in its native range and feared for its devastating effects on native diversity where the species is introduced. Lack of successful control in North America resulted in the initiation of a biological control program. We used a combination of field surveys and common garden experiments in Europe to study life history and ecology of a chloropid fly, Platycephala planifrons, to assess its potential as a biological control agent. The fly is widely distributed (in non-flooded sites) throughout Eurasia but attack rates are generally low (mean 5–10%; max. 29%). Adults emerge in late June and may live for several months. Females lay eggs at the base of Ph. australis shoots. First instar larvae of this stem-feeding fly overwinter in dormant below-ground shoots of Ph. australis and rapidly complete development in early spring. Larval feeding destroys the growing meristem of the shoot causing premature wilting and 60–70% reductions in shoot biomass production. Early season attack and considerable impact suggest that Pl. planifrons could be a potent biocontrol agent, if it can escape suppression by natural enemies in the introduced range. However, the generally low attack rates in its native range and its dependence on dry sites appear to make the species a “second-choice” candidate for potential release in North America.     

The suitability of non-target native mangroves for the survival and development of the lantana bug <Emphasis Type="Italic">Aconophora compressa</Emphasis>, an introduced weed biological control agent

Aconophora compressa (Hemiptera: Membracidae), a biological control agent introduced against the weed Lantana camara (Verbenaceae) in Australia, has since been observed on several non-target plant species, including native mangrove Avicennia marina (Acanthaceae). In this study we evaluated the suitability of two native mangroves, A. marina and Aegiceras corniculatum (Myrsinaceae), for the survival and development of A. compressa through no-choice field cage studies. The longevity of females was significantly higher on L. camara (57.7 ± 3.8 days) than on A. marina (43.3 ± 3.3 days) and A. corniculatum (45.7 ± 3.8 days). The proportion of females laying eggs was highest on L. camara (72%) followed by A. marina (36%) and A. corniculatum (17%). More egg batches per female were laid on L. camara than on A. marina and A. corniculatum. Though more nymphs per shoot emerged on L. camara (29.9 ± 2.8) than on A. marina (13 ± 4.8) and A. corniculatum (10 ± 5.3), the number of nymphs that developed through to adults was not significantly different. The duration of nymphal development was longer on A. marina (67 ± 5.8 days) than on L. camara (48 ± 4 days) and A. corniculatum (43 ± 4.6 days). The results, which are in contrast to those from previous glasshouse and quarantine trials, provide evidence that A. compressa adults can survive, lay eggs and complete nymphal development on the two non-target native mangroves in the field under no-choice condition.     

A new forest pest in Europe: a review of Emerald ash borer (Agrilus planipennis) invasion

In this publication, we review the biology, ecology, invasion history, impacts and management options of Emerald ash borer (EAB) Agrilus plannipennis, with a particular focus on its invasion in Europe. Agrilus planipennis (EAB) is a wood‐boring beetle native to East Asia. Having caused massive damages on ash species in North America in the last decades, it was first recorded in Europe in 2003 in Russia (Moscow). All ash (Fraxinus) species native to Europe and North America are known to be susceptible to EAB attacks, which cause high tree mortality even among formerly healthy trees. Recorded expansion rates are between 2.5 and 80 km/year in North America and between 13 and 41 km/year in European Russia. Given current expansion rates, EAB is expected to reach Central Europe within 15–20 years. A combination of mechanical, biological and chemical control and phytosanitary measures may reduce its impact, which nevertheless most likely will be substantial. There is an urgent need to identify native enemies in Europe, to test suitable biocontrol agents and to develop early detection and management measures. Although it is obvious that EAB will become a major pest in Europe, early and dedicated response will likely be able to reduce the level of ash mortality, and thus improve the opportunity for long‐term survival of ash as an important component in European forests.     

Nosema maddoxi sp. nov. (Microsporidia,Nosematidae), a Widespread Pathogen of the Green Stink Bug Chinavia hilaris (Say) and the Brown Marmorated Stink Bug Halyomorpha halys (Stål)

We describe a unique microsporidian species that infects the green stink bug, Chinavia hilaris; the brown marmorated stink bug, Halyomorpha halys; the brown stink bug, Euschistus servus; and the dusky stink bug, Euschistus tristigmus. All life stages are unikaryotic, but analysis of the consensus small subunit region of the ribosomal gene places this microsporidium in the genus Nosema, which historically has been characterized by diplokaryotic life stages. It is also characterized by having the reversed arrangement of the ribosomal gene (LSU –ITS‐ SSU) found in species within the “true Nosema” clade. This microsporidium is apparently Holarctic in distribution. It is present in H. halys both where it is native in Asia and where it is invasive in North America, as well as in samples of North American native C. hilaris collected prior to the introduction of H. halys from Asia. Prevalence in H. halys from mid‐Atlantic, North America in 2015–2016 ranged from 0.0% to 28.3%, while prevalence in C. hilaris collected in Illinois in 1970–1972 ranged from 14.3% to 58.8%. Oral infectivity and pathogenicity were confirmed in H. halys and C. hilaris. Morphological, ultrastructural, and ecological features of the microsporidium, together with a molecular phylogeny, establish a new species named Nosema maddoxi sp. nov.     

Biology and host specificity of Ceutorhynchus scrobicollis (Curculionidae; Coleoptera), a root-crown mining weevil proposed as biological control agent against Alliaria petiolata in North America

Ceutorhynchus scrobicollis is a root-crown mining weevil proposed for release as biological control agent of Alliaria petiolata (Brassicaceae, Thlaspideae), a European biennial herb, currently invading temperate North America. Using a combination of laboratory, common garden and fieldwork we studied biology, ecology and host range of C. scrobicollis, a univoltine species that oviposits and develops in A. petiolata rosettes in fall and spring. Individual C. scrobicollis can be long-lived (>2 years) and females show a second oviposition period. Weevils did not attack any of 31 test plant species outside the Brassicaceae. Within the Brassicaceae, five species allowed complete larval development under no-choice conditions. In subsequent choice tests, three of these five species (Nasturtium officinale, Peltaria alliacea and Thlaspi arvense; which are of European origin) were attacked. North American Rorippa sinuata was the only native species to be attacked by C. scrobicollis and only under no-choice conditions. Results of subsequent impact experiments showed that C. scrobicollis attack changed plant architecture but had no effect on overall plant vigour and reproductive output of R. sinuata, suggesting lack of impact on demography or population dynamics. A petition for field release of C. scrobicollis in North America has been submitted.     

Twenty‐five years after: post‐introduction association of Mecinus janthinus s.l. with invasive host toadflaxes Linaria vulgaris and Linaria dalmatica in North America

Linaria vulgaris, common or yellow toadflax, and Linaria dalmatica, Dalmatian toadflax (Plantaginaceae), are Eurasian perennial forbs invasive throughout temperate North America. These Linaria species have been the targets of classical biological control programmes in Canada and the USA since the 1960s. The first effective toadflax biological control agent, the stem‐mining weevil Mecinus janthinus (Coleoptera: Curculionidae) was introduced from Europe in the 1990s. This weevil has become established on L. dalmatica and L. vulgaris in both countries, although it has shown greater success in controlling the former toadflax species. Genetic and ecological studies of native range M. janthinus populations revealed that weevils previously identified as a single species in fact include two cryptic species, now recognised as M. janthinus, associated with yellow toadflax, and the recently confirmed species Mecinus janthiniformis, associated with Dalmatian toadflax. The results of a comprehensive study characterising haplotype identities, distributions and frequencies within M. janthinus s.l. native range source populations were compared to those populations currently established in the USA and Canada. The presence of both Mecinus species in North America was confirmed, and revealed with a few exceptions a high and consistent level of host fidelity throughout the adopted and native ranges. Genetic analysis based on mitochondrial cytochrome oxidase subunit II gene (mtCOII) defined the origin and records the subsequent North American establishment, by haplotype, of the European founder populations of M. janthinus (northern Switzerland and southern Germany) and M. janthiniformis (southern Macedonia), and provided population genetic indices for the studied populations. This analysis together with existing North American shipment receipt, release and rearing records elucidates probable redistribution routes and sources of both weevil species from initially released and established adopted range populations.     

Global invasion by <Emphasis Type="Italic">Anthidium manicatum</Emphasis> (Linnaeus) (Hymenoptera: Megachilidae): assessing potential distribution in North America and beyond

The wool carder bee, Anthidium manicatum, is the most widely distributed unmanaged bee in the world. It was unintentionally introduced to North America in the late 1960s from Europe, and subsequently, into South America, New Zealand and the Canary Islands. We provide information on the local distribution, seasonal abundance and sex ratio of A. manicatum from samples collected in an intensive two-year survey across Utah, USA. Anthidium manicatum was detected in 10 of the 29 Utah counties, largely in urban and suburban settings. Combining presence-only and MaxEnt background data from literature, museum databases and new records from Utah, we constructed three species distribution models to examine the potential distribution of A. manicatum in its native Eurasian range as well as invaded ranges of North and South America. The A. manicatum model based on locality and background data from the species’ native range predicted 50% of the invasive records associated with high habitat suitability (HS ≥ 0.90). The invasive North American model predicted a much broader distribution of A. manicatum (214% increase); whereas, the South American model predicted a narrower distribution (88% decrease). The poor predictive power of the latter model in estimating suitable habitats in the invasive South American range of A. manicatum suggests that the bee may still be limited by the bioclimatic constraints associated with a novel environment. Estimates of niche similarity (D) between the native and invasive models find that the North America bioclimatic niche is more similar to the bioclimatic niche of the native model (D = 0.78), whereas the bioclimatic niche of the South America invasion is relatively dissimilar (D = 0.69). We discuss the naturalization of A. manicatum in North America, possibly through punctuated dispersal, the probability of suitable habitats across the globe and the synanthropy exhibited by this invasive species.     

Dreyfusia nordmannianae in Northern and Central Europe: potential for biological control and comments on its taxonomy

The silver fir woolly adelgid, Dreyfusia nordmannianae, is the most severe pest occurring on Abies nordmanniana in Central and Northern Europe. The adelgid is particularly damaging to trees in Christmas tree plantations. Dreyfusia nordmannianae is native to the Caucasus region and alien to Europe, where its natural enemy complex is less diverse compared to its area of origin. Mitochondrial and nuclear DNA sequence data from the samples of D. nordmannianae collected in its native range and Europe and from the samples of Dreyfusia piceae and Dreyfusia prelli collected in Europe and North America were examined for phylogenetic structure. There was no evidence of differentiation, suggesting that these Dreyfusia species have recently diverged or require taxonomic revision. All existing published and unpublished reports on natural enemies of D. nordmannianae in its place of origin were reviewed, with the purpose of selecting agents for classical biological control in Europe. The literature review suggested that the most promising agent was the Chamaemyiidae fly, Leucopis hennigrata. A new survey in D. nordmannianae's area of origin – Turkey, Georgia and Russia – showed that L. hennigrata was present in all localities. It was particularly abundant in Turkey, where its impact on populations of D. nordmannianae appears to be high. Its use as a biological control agent is discussed, as well as other biological control strategies.     

Similar experiment – different conclusion; a response to the recent proposition of Septoria lepidii as a potentially important biological control agent for weedy species of Lepidium in North America

Following preliminary host-specificity testing Septori lepidii Desm. was recently proposed as a potentially important biological control agent for invasive Lepidium species in North America. Conversely, a previous, unpublished assessment of S. lepidii for biocontrol of Lepidium latifolium found the pathogen to be unsuitable due to infection of the American native Lepidium huberi.     

Biological control of Rhamnus cathartica: is it feasible? A review of work done in 2002–2012

Rhamnus cathartica (common buckthorn) is a shrub (or small tree) of Eurasian origin, which has become invasive in North America. Internal feeders and sap suckers were prioritized for biological control from over 30 specialized insects identified from the target plant in its native European range. Five leaf‐feeding moths were also considered for further investigations. Field observations and preliminary host range tests with the stem‐boring beetle Oberea pedemontana, the root‐boring moth Synanthedon stomoxiformis, the shoot‐tip‐boring moth Sorhagenia janiszewskae and the leaf‐feeding moths Ancylis apicella, A. unculana, Triphosa dubitata, Philereme transversata and P. vetulata confirmed that all of these species were lacking host specificity in no‐choice conditions. Choice oviposition tests carried out with most of the prioritized species to assess their ecological host range yielded unreliable results. Three psyllids, Trichochermes walkeri, Cacopsylla rhamnicolla and Trioza rhamni are promising in terms of host specificity, but are infected with the plant disease ‘Candidatus Phytoplasma rhamni’. Fruit‐ or seed‐feeding insects may present the best potential for biological control of buckthorn in directly reducing seed set and thus seedling establishment. However, it was not possible to obtain adult fruiting trees of native North American Rhamnus species for testing. It is concluded that there are no promising arthropod agents based on what is known to date. Pathogens could offer new opportunities for biological control of R. cathartica in North America.     

Laboratory host range of Austromusotima camptozonale (Lepidoptera: Crambidae), a potential biological control agent of Old World climbing fern,Lygodium microphyllum (Lygodiaceae)

Old World climbing fern, Lygodium microphyllum, is one of the most serious invasive weeds impacting south Florida and development of biological control is crucial for sustainable management. Larvae of a small moth, Austromusotima camptozonale, were discovered defoliating L. microphyllum in Australia. Preliminary testing suggested this moth was a Lygodium specialist. Laboratory host range testing was conducted on 65 species of test plants, from 31 families, comprising seven Lygodium species, four close relatives, 45 other species of ferns and fern allies, eight agricultural crops and one gymnosperm species plus the primary host L. microphyllum. Significant oviposition occurred only on other species of Lygodium. No eggs were laid on the agricultural crops, or about half the species of non-Lygodium ferns and fern allies tested. Oviposition on the other non-Lygodium ferns was very low, except on Anemia adiantifolia and Blechnum serrulatum, which received modest egg loads, but did not support development to adult. Larval feeding was low to non-existent on all the non-Lygodium species. Larvae developed to adult only on the native, American climbing fern L. palmatum, and to a lesser extent on L. japonicum. Lygodium japonicum is a naturalized invasive weed in the United States. Colonies of A. camptozonale were unable to persist on L. palmatum and died out in two to seven generations. Freezing winter temperatures in states where L. palmatum occurs would be lethal to A. camptozonale. It was concluded that A. camptozonale would pose no threat to native or cultivated plants in North America or the Caribbean and should be considered as a weed biological control agent against L. microphyllum.