Hosts and impacts of elongate hemlock scale (Hemiptera: Diaspididae): A critical review.

Fiorinia externa Ferris, elongate hemlock scale, was inadvertently introduced to North America from Japan. This insect is particularly problematic on hemlock, Tsuga spp., though it has been reported in association with several other conifers. The evidence that other conifers might be hosts, capable of supporting growing populations of the insect, has not been previously reviewed. Our review confirms that F. externa is an oligophagous pest of members of Pinaceae. Although species of Cupressaceae and Taxaceae have been reported as hosts of F. externa, they seem unable to support population growth of this pest. Evidence of the tree-killing potential of the insect, even on suitable hosts, is remarkably scant. The degree of pest risk posed by F. externa with respect to tree mortality in areas beyond the geographic range of hemlock seems modest, but uncertain.

Pest population dynamics are related to a continental overwintering gradient.

Overwintering success is an important determinant of arthropod populations that must be considered as climate change continues to influence the spatiotemporal population dynamics of agricultural pests. Using a long-term monitoring database and biologically relevant overwintering zones, we modeled the annual and seasonal population dynamics of a common pest, Helicoverpa zea (Boddie), based on three overwintering suitability zones throughout North America using four decades of soil temperatures: the southern range (able to persist through winter), transitional zone (uncertain overwintering survivorship), and northern limits (unable to survive winter). Our model indicates H. zea population dynamics are hierarchically structured with continental-level effects that are partitioned into three geographic zones. Seasonal populations were initially detected in the southern range, where they experienced multiple large population peaks. All three zones experienced a final peak between late July (southern range) and mid-August to mid-September (transitional zone and northern limits). The southern range expanded by 3% since 1981 and is projected to increase by twofold by 2099 but the areas of other zones are expected to decrease in the future. These changes suggest larger populations may persist at higher latitudes in the future due to reduced low-temperature lethal events during winter. Because H. zea is a highly migratory pest, predicting when populations accumulate in one region can inform synchronous or lagged population development in other regions. We show the value of combining long-term datasets, remotely sensed data, and laboratory findings to inform forecasting of insect pests.

Potential displacement of the native Tenuisvalvae notata by the invasive Cryptolaemus montrouzieri in South America suggested by differences in climate suitability.

Tenuisvalvae notata (Mulsant) (Coccinellidae) is a predatory ladybird beetle native to South America. It specializes in mealybugs prey (Pseudococcidae), but relatively little is known about its ecology. In contrast, the ladybird beetle Cryptolaemus montrouzieri Mulsant (Coccinellidae) is indigenous to Australia and has been introduced to many countries worldwide including Brazil for biological control of mealybugs. The potential impacts of these introductions to native coccinellids have rarely been considered. The software CLIMEX estimated the climate suitability for both species as reflected in the Ecoclimatic Index (EI). Much of South America, Africa, and Australia can be considered climatically suitable for both species, but in most cases, the climate is considerably more favorable for C. montrouzieri than T. notata, especially in South America. The CLIMEX model also suggests seasonal differences in growth conditions (e.g. rainfall and temperature) that could affect the phenology of both species. These models suggest that few locations in South America would be expected to provide T. notata climatic refugia from C. montrouzieri. Although other ecological factors will also be important, such as prey availability, this analysis suggests a strong potential for displacement of a native coccinellid throughout most of its range as a consequence of the invasion by an alien competitor.

Balancing the need for seed against invasive species risks in prairie habitat restorations.

Adequate diversity and abundance of native seed for large-scale grassland restorations often require commercially produced seed from distant sources. However, as sourcing distance increases, the likelihood of inadvertent introduction of multiple novel, non-native weed species as seed contaminants also increases. We created a model to determine an "optimal maximum distance" that would maximize availability of native prairie seed from commercial sources while minimizing the risk of novel invasive weeds via contamination. The model focused on the central portion of the Level II temperate prairie ecoregion in the Midwest US. The median optimal maximum distance from which to source seed was 272 km (169 miles). In addition, we weighted the model to address potential concerns from restoration practitioners: 1. sourcing seed via a facilitated migration strategy (i.e., direct movement of species from areas south of a given restoration site to assist species' range expansion) to account for warming due to climate change; and 2. emphasizing non-native, exotic species with a federal mandate to control. Weighting the model for climate change increased the median optimal maximum distance to 398 km (247 miles), but this was not statistically different from the distance calculated without taking sourcing for climate adaptation into account. Weighting the model for federally mandated exotic species increased the median optimal maximum distance only slightly to 293 km (182 miles), so practitioners may not need to adjust their sourcing strategy, compared to the original model. This decision framework highlights some potential inadvertent consequences from species translocations and provides insight on how to balance needs for prairie seed against those risks.

Sublethal effects of subzero temperatures on the light brown apple moth, Epiphyas postvittana: fitness costs in response to partial freezing.

Population responses to environmental extremes often dictate the bounds to species' distributions. However, population dynamics at, or near, those range limits may also be affected by sublethal effects. We exposed late instars and pupae of an invasive leafroller, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), to cold temperatures and measured the effects of exposure on subsequent survivorship, development, and reproduction. Cold temperature was applied as acute exposure to -10 °C (a low, but not immediately lethal temperature for this species) or the onset of freezing (the peak of the supercooling point exotherm). Survival was defined as the ability to successfully eclose as an adult. We measured immature development times, pupal mass, and adult longevity as proxies of fitness in survivors. Additionally, surviving insects were mated with individuals that had not been exposed to cold to measure fertility. There was no difference between the proportion of larvae or pupae that survived acute exposure to -10 °C and those exposed to the control temperature. Approximately 17% of larvae and 8% of pupae survived brief periods with internal ice formation and continued development to become reproductively viable adults. Importantly, surviving the onset of freezing came with significant fitness costs but not to exposure to -10 °C; most insects that survived partial freezing had lower fertility and shorter adult lifespans than either the -10 °C or control group. These results are discussed within the context of forecasting invasive insect distributions.

Effects of Diapause on Halyomorpha halys (Hemiptera: Pentatomidae) Cold Tolerance.

Diapause and cold tolerance can profoundly affect the distribution and activity of temperate insects. Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), an alien invasive species from Asia, enters a winter dormancy in response to environmental cues. We investigated the nature of this dormancy and its effects on H. halys cold tolerance, as measured by supercooling points, lower lethal temperatures, and overwintering field mortality. Dormancy was induced by rearing individuals in the laboratory or under field conditions. We confirmed H. halys dormancy to be a state of diapause and not quiescence, and the life stage sensitive to diapause-inducing cues is between the second and fifth instar. In the laboratory, supercooling points of diapausing adults reached significantly lower temperatures than nondiapausing adults, but only when given enough time after imaginal ecdysis. Supercooling points of diapausing adults in overwintering microhabitats also decreased over time. Diapause increased adult survival after acute cold exposure in the laboratory and prolonged cold exposure in the field. Following diapause induction in the laboratory, changes to temperature and photoperiod had no significant effect on lower lethal temperatures and changes to photoperiod had no effect on supercooling points. Additionally, induction of diapause in the laboratory did not result in significantly different cold tolerance than natural field induction of diapause. This work demonstrates that H. halys diapause confers greater cold tolerance than a nondiapausing state and likely improves the probability of successful overwintering in some temperate climates. Hence, knowledge of diapause status could be used to refine forecasts of H. halys overwintering field mortality.

Effects of Adult Feeding and Overwintering Conditions on Energy Reserves and Flight Performance of Emerald Ash Borer (Coleoptera: Buprestidae).

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive beetle from Asia, spreads through human-mediated movement and active flight. The effects of adult feeding and overwintering conditions on A. planipennis energy reserves (e.g., lipid, glycogen, and sugars) and flight are poorly understood. We conjectured that the potential energetic demands associated with the production of cryoprotectants might affect dispersal capacity and partially explain slower spread of A. planipennis in Minnesota than in the other states. Two studies sought to measure the effects of adult feeding on lipid content and flight capacity. Adult A. planipennis were fed shamel ash, Fraxinus uhdei Wenzig, leaves for 0-20 d after emergence, and half were flown on a custom flight mill for 24 h, before being frozen for comparative lipid analysis with a control group. The second study compared the effects of adult feeding on energy reserves and flight capacity of A. planipennis that were originally from St. Paul, Minnesota but overwintered in infested logs placed in Grand Rapids, Minnesota (low winter temperature, -34°C) or St. Paul, Minnesota (-26.3°C). Live adults consumed foliage at a constant rate, but lipid content (percentage of fresh mass) did not change with increases in feeding or flight. Adult glycogen content declined with flight and increased only slightly with feeding. Overwintering location affected survival rates but not energy reserves or flight capacity. These results suggest that the flight capacity of A. planipennis is largely determined before emergence, with no differences in energy reserves after cryoprotectant investment.

Visitor Preferences for Visual Changes in Bark Beetle-Impacted Forest Recreation Settings in the United States and Germany.

Extensive outbreaks of tree-killing insects are increasing across forests in Europe and North America due to climate change and other factors. Yet, little recent research examines visitor response to visual changes in conifer forest recreation settings resulting from forest insect infestations, how visitors weigh trade-offs between physical and social forest environment factors, or how visitor preferences might differ by nationality. This study explored forest visitor preferences with a discrete choice experiment that photographically simulated conifer forest stands with varying levels of bark beetle outbreaks, forest and visitor management practices, and visitor use levels and compositions. On-site surveys were conducted with visitors to State Forest State Park in Colorado (n = 200), Lake Bemidji State Park in Minnesota (n = 228), and Harz National Park in Germany (n = 208). Results revealed that the condition of the immediate forest surrounding was the most important variable influencing visitors' landscape preferences. Visitors preferred healthy mature forest stands and disliked forests with substantial dead wood. The number of visitors was the most important social factor influencing visitor landscape preferences. Differences in the influence of physical and social factors on visual preferences existed between study sites. Findings suggest that both visual forest conditions and visitor use management are important concerns in addressing landscape preferences for beetle-impacted forest recreation areas.

Cold Tolerance of Pityophthorus juglandis (Coleoptera: Scolytidae) From Northern California.

Winter survivorship of insects is determined by a combination of physiological, behavioral, and microhabitat characteristics. We characterized the cold tolerance of the walnut twig beetle, Pityophthorus juglandis Blackman, a domestic alien invasive bark beetle that vectors a phytopathogenic fungus. The beetle and fungus cause thousand cankers disease in species of Juglans and Pterocarya. The disease is spreading in the United States of America (USA) and Italy. Contact thermocouple thermometry was used to measure the supercooling points of adults and larvae and lower lethal temperatures of adults from a population from northern California. Supercooling points ranged from -12.2 °C to - 25.0 °C for adults and -13.6 °C to - 23.5 °C for larvae; lower lethal temperatures of adults ranged from -14 °C to - 23 °C. We found seasonal changes in adult supercooling points in fall, winter, and spring. The supercooling point for males was 0.5 °C colder than for females over all months and 1 °C colder in the winter than in other seasons. The cold-tolerance strategy shifted in P. juglandis adults from freeze intolerance (December 2013 and January 2014) to partial freeze tolerance (February 2014). An intermediate level of cold tolerance with a plastic response to cold partially explains survival of P. juglandis outside of its native range in the southwestern USA. In addition, we characterized the relationship between minimum air temperatures and minimum phloem temperatures in two Juglans spp. in northern California and Colorado and characterized portions of the native geographic range of eastern black walnut, J. nigra L., that may be too cold currently for this insect to persist.

Colonization behaviors of mountain pine beetle on novel hosts: Implications for range expansion into northeastern North America.

As climates change, thermal limits may no longer constrain some native herbivores within their historical ranges. The mountain pine beetle, Dendroctonus ponderosae Hopkins, is a tree-killing bark beetle native to western North America that is currently expanding its range. Continued eastward expansion through the newly invaded and novel jack pine (Pinus banksiana Lamb.) trees of the Canadian boreal forest could result in exposure of several species of novel potential host pines common in northeastern North America to this oligophagous herbivore. Due to the tightly co-evolved relationship between mountain pine beetle and western pine hosts, in which the insect utilizes the defensive chemistry of the host to stimulate mass attacks, we hypothesized that lack of co-evolutionary association would affect the host attraction and acceptance behaviors of this insect among novel hosts, particularly those with little known historical association with an aggressive stem-infesting insect. We studied how beetle behavior differed among the various stages of colonization on newly cut logs of four novel potential pine host species; jack, red (P. resinosa Ait.), eastern white (P. strobus L.) and Scots (P. sylvestris L.) pines, as well as two historical hosts, ponderosa (P. ponderosa Dougl. ex. Laws. var. scopulorum Engelm.) and lodgepole (P. contorta Dougl. var. latifolia Engelm.) pines. Overall, we found that beetle colonization behaviors at each stage in the colonization process differ between pine hosts, likely due to differing chemical and physical bark traits. Pines without co-evolved constitutive defenses against mountain pine beetle exhibited reduced amounts of defensive monoterpenoid chemicals; however, such patterns also reduced beetle attraction and colonization. Neither chemical nor physical defenses fully defended trees against the various stages of host procurement that can result in tree colonization and death.

Flight Capacity of the Walnut Twig Beetle (Coleoptera: Scolytidae) on a Laboratory Flight Mill.

The walnut twig beetle, Pityophthorus juglandis Blackman, and associated fungus Geosmithia morbida Kolarík, Freeland, Utley, & Tisserat constitute the insect-fungal complex that causes thousand cankers disease in walnut, Juglans spp., and wingnut, Pterocarya spp. Thousand cankers disease is responsible for the decline of Juglans species throughout the western United States and more recently, the eastern United States and northern Italy. We examined the flight capacity of P. juglandis over 24-h trials on a flight mill in the laboratory. The maximum total flight distance observed was ∼3.6 km in 24 h; however, the mean and median distances flown by beetles that initiated flight were ∼372 m and ∼158 m, respectively. Beetles flew for 34 min on average within a 24-h flight trial. Male and female flight capacities were similar, even though males were larger than females (0.64 vs. 0.57 mm pronotal width). Age postemergence had no effect on flight distance, flight time, or mean flight velocity. The propensity to fly, however, decreased with age. We integrated results of flight distance with propensity to fly as beetles aged in a Monte Carlo simulation to estimate the maximum dispersal capacity over 5 d, assuming no mortality. Only 1% of the insects would be expected to fly >2 km, whereas one-third of the insects were estimated to fly <100 m. These results suggest that nascent establishments remain relatively localized without anthropogenic transport or wind-aided dispersal, which has implications for management and sampling of this hardwood pest.

Host-mediated shift in the cold tolerance of an invasive insect.

While many insects cannot survive the formation of ice within their bodies, a few species can. On the evolutionary continuum from freeze-intolerant (i.e., freeze-avoidant) to freeze-tolerant insects, intermediates likely exist that can withstand some ice formation, but not enough to be considered fully freeze tolerant. Theory suggests that freeze tolerance should be favored over freeze avoidance among individuals that have low relative fitness before exposure to cold. For phytophagous insects, numerous studies have shown that host (or nutrition) can affect fitness and cold-tolerance strategy, respectively, but no research has investigated whether changes in fitness caused by different hosts of polyphagous species could lead to systematic changes in cold-tolerance strategy. We tested this relationship with the invasive, polyphagous moth, Epiphyas postvittana (Walker). Host affected components of fitness, such as larval survivorship rates, pupal mass, and immature developmental times. Host species also caused a dramatic change in survival of late-instar larvae after the onset of freezing-from less than 8% to nearly 80%. The degree of survival after the onset of freezing was inversely correlated with components of fitness in the absence of cold exposure. Our research is the first empirical evidence of an evolutionary mechanism that may drive changes in cold-tolerance strategies. Additionally, characterizing the effects of host plants on insect cold tolerance will enhance forecasts of invasive species dynamics, especially under climate change.

Cold Tolerance of Halyomorpha halys (Hemiptera: Pentatomidae) Across Geographic and Temporal Scales.

The brown marmorated stink bug, Halyomorpha halys (Stål), is native to eastern Asia and is presently invading North America. Little is known about the exposure to and effects of winter temperatures in newly invaded regions on H. halys The overwintering habitats that this species utilizes vary greatly in their thermal buffering capacity. They naturally overwinter in aggregations beneath loose bark on trees and in cliff outcroppings, but will also commonly aggregate in buildings. Effects of cold temperatures such as mortality and freezing have yet to be quantified in the invading population. We report that H. halys is chill intolerant (i.e., dies before reaching its freezing point), and that the degree of cold tolerance of populations in North America differs by season, sex, and acclimation location. The mean winter supercooling point (± SEM) of individuals acclimated in Minnesota was -17.06 °C ± 0.13 and in Virginia was -13.90 °C ± 0.09. By using laboratory assays of lower lethal temperatures and ambient air temperature records, we accurately forecasted mortality for field experiments in Minnesota and Virginia. Temperature refugia provided by human-built structures are likely crucial for overwintering survival during atypically cold winters and possibly contribute to the northern geographic range expansion of this economically damaging insect in the temperate climates of North America.

Densities of Agrilus auroguttatus and Other Borers in California and Arizona Oaks.

We investigated within-tree population density of a new invasive species in southern California, the goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), with respect to host species and the community of other borers present. We measured emergence hole densities of A. auroguttatus and other borers on the lower stem (bole) of naïve oaks at 18 sites in southern California and on co-evolved oaks at seven sites in southeastern Arizona. We sampled recently dead oaks in an effort to quantify the community of primary and secondary borers associated with mortality-species that were likely to interact with A. auroguttatus. Red oaks (Section Lobatae) produced greater densities of A. auroguttatus than white oaks (Section Quercus). On red oaks, A. auroguttatus significantly outnumbered native borers in California (mean ± SE of 9.6 ± 0.7 versus 4.5 ± 0.6 emergence holes per 0.09 m² of bark surface), yet this was not the case in Arizona (0.9 ± 0.2 versus 1.1 ± 0.2 emergence holes per 0.09 m²). In California, a species that is taxonomically intermediate between red and white oaks, Quercus chrysolepis (Section Protobalanus), exhibited similar A. auroguttatus emergence densities compared with a co-occurring red oak, Q. kelloggii. As an invasive species in California, A. auroguttatus may affect the community of native borers (mainly Buprestidae and Cerambycidae) that feed on the lower boles of oaks, although it remains unclear whether its impact will be positive or negative.

Assessing the cost of an invasive forest pathogen: a case study with oak wilt.

Economic assessment of damage caused by invasive alien species provides useful information to consider when determining whether management programs should be established, modified, or discontinued. We estimate the baseline economic damage from an invasive alien pathogen, Ceratocystis fagacearum, a fungus that causes oak wilt, which is a significant disease of oaks (Quercus spp.) in the central United States. We focus on Anoka County, Minnesota, a 1,156 km(2) mostly urban county in the Minneapolis-Saint Paul metropolitan region. We develop a landscape-level model of oak wilt spread that accounts for underground and overland pathogen transmission. We predict the economic damage of tree mortality from oak wilt spread in the absence of management during the period 2007-2016. Our metric of economic damage is removal cost, which is one component of the total economic loss from tree mortality. We estimate that Anoka County has 5.92 million oak trees and 885 active oak wilt pockets covering 5.47 km(2) in 2007. The likelihood that landowners remove infected oaks varies by land use and ranges from 86% on developed land to 57% on forest land. Over the next decade, depending on the rates of oak wilt pocket establishment and expansion, 76-266 thousand trees will be infected with discounted removal cost of $18-60 million. Although our predictions of removal costs are substantial, they are lower bounds on the total economic loss from tree mortality because we do not estimate economic losses from reduced services and increased hazards. Our predictions suggest that there are significant economic benefits, in terms of damage reduction, from preventing new pocket establishment or slowing the radial growth of existing pockets.

Boring in response to bark and phloem extracts from North American trees does not explain host acceptance behavior of Orthotomicus erosus (Coleoptera: Scolytidae).

When invasive herbivorous insects encounter novel plant species, they must determine whether the novel plants are hosts. The Mediterranean pine engraver, Orthotomicus erosus (Wollaston), an exotic bark beetle poised to expand its range in North America, accepts hosts after contacting the bark. To test the hypothesis that O. erosus accepts hosts on the basis of gustatory cues, we prepared bark and phloem extracts from logs of four North American tree species that we had used in previous host acceptance experiments. Water, methanol, and hexane extracts of red pine, tamarack, balsam fir, and paper birch were presented alone and in combination on a neutral filter paper substrate in a section of a plastic drinking straw. Boring behavior in response to the three-extract combinations differed from the pattern of acceptance previously observed among species when the beetles were in contact with the bark surface. Only the aqueous extracts of tamarack, Larix laricina, increased the initiation and the extent of boring by O. erosus on the filter paper substrate. We conclude that the effects of extracted chemicals do not match the behavior of the beetles observed when penetrating excised bark and phloem discs, indicating that host selection by O. erosus may not be predictable from bark and phloem chemistry alone. Instead, host acceptance may be determined by nongustatory stimuli or by a combination of stimuli including gustatory and nongustatory cues.

Invasions by Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in the Western Hemisphere: implications for South America.

The multicolored Asian lady beetle, Harmonia axyridis (Pallas), native to Asia, has recently been detected in South America after successfully invading North America and Europe. This coccinellid is a voracious predator; therefore, it is popular and effective in biological control. Unfortunately, H. axyridis also has associated adverse impacts (i.e., as a household pest, pest of fruit production, and threat to non-target organisms). To predict the potential geographic extent of impacts of H. axyridis in South America we review the history of its invasion in the Western Hemisphere and address various factors critical to the future invasion (i.e., arrival, establishment, and spread) of new areas of South America. The likelihood of continued introductions (i.e., arrival) of H. axyridis to South America seems high, due to its popularity as a biological control agent and through accidental introductions. Establishment also seems likely in broad regions of South America. Climate matching with the native range suggested that much of southern South America may be suitable for establishment. In contrast, habitat matching with the native range suggested that northern South America may be more suitable. In addition, prey availability should not limit establishment of this predator. Once established, H. axyridis seems likely to spread by flight and human-assisted means. Overall, the invasion of H. axyridis over broad areas in South America seems likely.

Invasions by Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in the Western Hemisphere: implications for South America

The multicolored Asian lady beetle, Harmonia axyridis (Pallas), native to Asia, has recently been detected in South America after successfully invading North America and Europe. This coccinellid is a voracious predator; therefore, it is popular and effective in biological control. Unfortunately, H. axyridis also has associated adverse impacts (i.e., as a household pest, pest of fruit production, and threat to non-target organisms). To predict the potential geographic extent of impacts of H. axyridis in South America we review the history of its invasion in the Western Hemisphere and address various factors critical to the future invasion (i.e., arrival, establishment, and spread) of new areas of South America. The likelihood of continued introductions (i.e., arrival) of H. axyridis to South America seems high, due to its popularity as a biological control agent and through accidental introductions. Establishment also seems likely in broad regions of South America. Climate matching with the native range suggested that much of southern South America may be suitable for establishment. In contrast, habitat matching with the native range suggested that northern South America may be more suitable. In addition, prey availability should not limit establishment of this predator. Once established, H. axyridis seems likely to spread by flight and human-assisted means. Overall, the invasion of H. axyridis over broad areas in South America seems likely.

A joaninha, Harmonia axyridis (Pallas), nativa da Asia, foi recentemente detectada na América do Sul depois de ter invadido a América do Norte e Europa. Essa joaninha é um predador voraz, e portanto, popular e eficaz no controle biológico. Infelizmente, H. axyridis também está relacionada a impactos nocivos (ex., como peste residencial e de frutas temperadas e ameaça a organismos não-alvos). Para fazer prever os possíveis impactos de H. axyridis na América do Sul, a história da sua invasão no Hemisfério Ocidental foi revisada e os vários fatores críticos para futuras invasões (isto é, chegada, estabelecimento e disseminação) em novas áreas da América do Sul foram discutidos. A possibilidade de introduções contínuas de H. axyridis na América do Sul parece alta devido a sua popularidade como agente de controle biológico e através de introduções acidentais. Seu estabelecimento também parece possível em extensas regiões da América do Sul. A similaridade climática com a região nativa sugere que o estabelecimento é possível na região sul da América do Sul. Porém, similaridade de hábitat com a região nativa sugere que o estabelecimento seja mais adequado na região norte da América do Sul. Além disso, a disponibilidade da presa não deve ser um fator limitante para o estabelecimento desse predador. Após o estabelecimento, H. axyridis pode se disseminar pelo próprio vôo e por meios associados ao homem. Concluindo, a invasão de H. axyridis em novas áreas do continente sul-americano é provável.

Strategies and statistics of sampling for rare individuals.

Diverse subdisciplines within entomology recognize the detection of rare individuals as the precursor to effective management of these individuals. Unfortunately, detection methods have often developed on a case-by-case basis, and advances in one subdiscipline have not carried over to similarly related fields. The biology of a particular organism will certainly affect sampling methods, but the underlying principles governing the power of a sampling strategy to detect rare individuals will apply across taxa. Our review of the sampling literature demonstrates the common problem of detecting rare individuals, reviews the fundamentals of probability theory as a foundation for any monitoring program, and discusses the inferences that can be drawn from samples, especially when resources limit sampling efforts. Particular emphasis is placed on binomial-, beta-binomial-, and hypergeometric-based sampling strategies as they pertain to quarantine inspections for exotic pests, veterinary/medical entomology, and insecticide resistance monitoring.