Realized Host Range Research Articles

Does the fundamental host range of Trissolcus japonicus match its realized host range in Europe?

Unintentional introduction of natural enemies has increased in recent years due to a massive rise in global trade and tourism. One such natural enemy is the Asian egg parasitoid Trissolcus japonicus, a promising agent for classical biological control of Halyomorpha halys. In Europe, adventive T. japonicus populations have been detected in Switzerland, Italy and Germany. Host specificity testing demonstrated that its fundamental host range is fairly broad; however, it is unclear whether spatial or temporal refuges reduce parasitism of non-target species in the field. To address this, the realized host range of T. japonicus was assessed over three years by exposing sentinel egg masses of H. halys and 18 non-target species and collecting naturally laid egg masses in Switzerland and Italy. In total, 15 of 18 non-target species were successfully parasitized by T. japonicus in the field, confirming its broad fundamental host range. However, most non-target species were less parasitized by T. japonicus than H. halys, profiting from either partial temporal or spatial refuges from parasitism. Species with an unusual life cycle and the same ecological niche as H. halys, such as Pentatoma rufipes, which was the most parasitized non-target species in both countries, potentially face an increased risk of parasitism. In contrast, beneficial non-target effects may occur for the invasive pest, Nezara viridula, which suffered high non-reproductive mortality induced by T. japonicus. In both cases, life table studies will be needed to determine the impact of non-target parasitism and the potential consequences at the population level.

From generalist to specialists: Variation in the host range and performance of anther-smut pathogens on Dianthus.

Determining the processes that drive the evolution of pathogen host range can inform our understanding of disease dynamics and the potential for host shifts. In natural populations, patterns of host range could be driven by genetically based differences in pathogen infectivity or ecological differences in host availability. In northwestern Italy, four reproductively isolated lineages of the fungal plant-pathogen Microbotryum have been shown to co-occur on several species in the genus Dianthus. We carried out cross-inoculation experiments to determine whether patterns of realized host range in these four lineages were driven by differences in infectivity and to test whether there was evidence of a trade-off between host range and within-host reproduction. We found strong concordance between field patterns of host range and pathogen infectivity on different Dianthus species using experimental inoculation, indicating that infection ability is a major driving force of host range. However, we found no evidence of a trade-off between the ability to infect a wider range of host species and spore production on a shared host.

Predicting non-target impacts.

Biocontrol of invasive alien weeds has produced great benefits, but concerns over undesirable impacts on non-target plants and/or indirect interactions between biocontrol agents and other biota impede the implementation of biocontrol in some countries. Although great strides have been made, continuing uncertainties predicting the realized host range of candidate agents is probably resulting in some being erroneously rejected due to overestimation of risk. Further refinement of host-range testing protocols is therefore desirable. Indirect interactions are inherently harder to predict, and the risk of both direct and indirect non-target impacts may change over time due to biocontrol agents evolving or expanding their range under climate change. Future research directions to better understand the risk of non-target impacts over time are discussed.

Host Range and Effects of Plant Species on Preference and Fitness of Tuta absoluta (Lepidoptera: Gelechiidae).

Tuta absoluta (Meyrick) is one of the most devastating pests of tomato in both open field and greenhouse production settings that significantly reduce yield and quality of the fruit. Being a new invasive pest, information on the pest bio-ecology, including host-rage and effects of plant species on its various fitness traits are virtually lacking. This study involved a field survey to establish the realized host range and laboratory experiments to evaluate the preference for, and fitness effects of T. absoluta on different plant species. In the field, T. absoluta was recovered from only four solanaceous plant species: tomato (Solanum lycopersicum L.), eggplant (S. melongena L.), black nightshade (S. nigrum L.), and gubbain (S. dubium Fr.). In choice and no-choice tests, tomato was the most preferred host of T. absoluta having the highest number of eggs deposited, followed by black nightshade, French bean Phaseolus vulgaris L., and chilli pepper Capsicum frutescens. Larvae developed quickest on tomato followed by black nightshade and French bean. The percentage survival to adulthood was significantly higher on tomato, compared to survival on black nightshade and French bean. Rearing of T. absoluta on tomato or black nightshade had no significant effect on pupal mass and length as well as adult size for both sexes. Fecundity and longevity, as well as oviposition preference of T. absoluta, were not affected by rearing on different plants. These findings confirm the oligophagous nature of the pest with high preference to tomato.

Tolerance of the eriophyid mite Aceria salsolae to UV-A light and implications for biological control of Russian thistle.

Aceria salsolae (Acari: Eriophyidae) is being evaluated as a candidate biological control agent of Russian thistle (Salsola tragus, Chenopodiaceae), a major invasive weed of rangelands and dryland crops in the western USA. Prior laboratory host range testing under artificial lighting indicated reproduction on non-native Bassia hyssopifolia and on a native plant, Suaeda calceoliformis. However, in field tests in the native range, mite populations released on these 'nontarget' plants remained low. We hypothesized that UV-A light, which can affect behavior of tetranychid mites, would affect populations of the eriophyid A. salsolae differently on the target and nontarget plant species, decreasing the mite's realized host range. Plants were infested with A. salsolae under lamps that emitted UV-A, along with broad-spectrum lighting, and the size of mite populations and plant growth was compared to infested plants exposed only to broad-spectrum light. Russian thistle supported 3- to 55-fold larger mite populations than nontarget plants regardless of UV-A treatment. UV-A exposure did not affect mite populations on Russian thistle or S. calceoliformis, whereas it increased populations 7-fold on B. hyssopifolia. Main stems on nontarget plants grew 2- to 6-fold faster than did Russian thistle under either light treatment. The two nontarget plants attained greater volume under the control light regime than UV-A, but Russian thistle was unaffected. Although Russian thistle was always the superior host, addition of UV-A light to the artificial lighting regime did not reduce the ability of A. salsolae to reproduce on the two nontarget species, suggesting that UV-B or other environmental factors may be more important in limiting mite populations in the field.

Assessing Probabilistic Risk Assessment Approaches for Insect Biological Control Introductions.

The introduction of biological control agents to new environments requires host specificity tests to estimate potential non-target impacts of a prospective agent. Currently, the approach is conservative, and is based on physiological host ranges determined under captive rearing conditions, without consideration for ecological factors that may influence realized host range. We use historical data and current field data from introduced parasitoids that attack an endemic Lepidoptera species in Hawaii to validate a probabilistic risk assessment (PRA) procedure for non-target impacts. We use data on known host range and habitat use in the place of origin of the parasitoids to determine whether contemporary levels of non-target parasitism could have been predicted using PRA. Our results show that reasonable predictions of potential non-target impacts may be made if comprehensive data are available from places of origin of biological control agents, but scant data produce poor predictions. Using apparent mortality data rather than marginal attack rate estimates in PRA resulted in over-estimates of predicted non-target impact. Incorporating ecological data into PRA models improved the predictive power of the risk assessments.

Host range expansion is density dependent.

The realized host range of herbivores is expected to increase with herbivore population density. Theory also predicts that trait similarity and phylogenetic relatedness between native and exotic plants is expected to increase the susceptibility of introduced plants to feeding by native herbivores. Whether the ability of native herbivores to extend their host range to introduced species is density dependent is still unknown. We addressed this question by monitoring pine processionary moth (PPM, Thaumetopoea pityocampa) attacks during nine consecutive years on 41 pine species (8 native and 33 introduced) planted in an arboretum. The survey encompassed latent and outbreak periods. A total of 28 pine species were attacked by PPM. There was no difference in the probability of attack between native and introduced pine species. Host range increased and was more phylogenetically clustered during outbreak than latent periods. When population density increased, PPM expanded its diet breadth by attacking introduced pine species that were closely related to native hosts. This study demonstrates the density dependence of host range expansion in a common pine herbivore. Importantly, it supports the idea that the degree of phylogenetic proximity between host species can be a better predictor of attacks than the introduction status, which may help to predict the outcomes of new plant-herbivore interactions.

Successes We May Not Have Had: A Retrospective Analysis of Selected Weed Biological Control Agents in the United States

AbstractIn this paper, we describe five successful classical biological weed control agents released in the United States. For each of the five arthropod species, we compared data from prerelease studies that experimentally predicted the agent's host range with data collected postrelease. In general, experimental host range data accurately predicted or overestimated risks to nontarget plants. We compare the five cases with insects recently denied for introduction in the United States and conclude that none of the discussed agents would likely be approved if they were petitioned today. Three agents would be rejected because they potentially could attack economic plants, and two because of potential attack on threatened or endangered plants. All five biocontrol agents have contributed significantly to the successful management of major weeds with no or minimal environmental risk. We believe that the United States may miss opportunities for sustainable and environmentally benign management of weeds using biological control if the regulatory framework only considers the risks of agents as potential plant pests and treats any host-range data regarding economic or threatened and endangered species as a binary decision (i.e., mandates rejection if there is any chance of feeding or development). As a way forward we propose the following: (1) the addition of risk and benefit analyses at the habitat level with a clear ranking of decision-making criteria as part of the U.S. Department of Agriculture Animal and Plant Health Inspection Service Technical Advisory Group's evaluation process of biocontrol agents; (2) recognition of the primacy of realized host range data for potential agents that considers the insect's host selection behavior instead of emphasizing fundamental host range data during release evaluations, and (3) development of formalized postrelease monitoring of target and nontarget species as part of the release permit. These recommendations may initially be advanced through reassessment of current policies but may in the longer term require the implementation of dedicated biocontrol legislation.

Factors affecting host range in a generalist seed pathogen of semi-arid shrublands

Generalist pathogens can exhibit differential success on different hosts, resulting in complex host range patterns. Several factors operate to reduce realized host range relative to potential host range, particularly under field conditions. We explored factors influencing host range of the naturally occurring generalist ascomycete grass seed pathogen Pyrenophora semeniperda .W e measured potential host range in laboratory experiments at high inoculum loads with 26 grass species, including the primary host Bromus tectorum, and developed models to predict susceptibility and tolerance based on host traits, including germination speed, seed hardness, seed size, and phylogenetic relations. We also examined pathogen and host density effects on infection and mortality. All species tested were at least somewhat susceptible to the pathogen at high inoculum loads, but both infection and mortality varied widely. Species more closely related to the original host (B. tectorum) were more susceptible to infection, whereas species with slower germination were less tolerant and therefore more likely to suffer mortality. Infection and mortality were sharply reduced as inoculum load was reduced. Intermediate loads had major negative impacts on dormant B. tectorum seeds but generally minimal effects on native species. In addition, field seed bank studies determined that P. semeniperda rarely exploits native grass species as hosts. This marked reduction in realized host range relative to potential host range indicates that laboratory host range studies are potentially a poor predictor of either the current or possible future realized host range for wildland plant pathogens.

Constraints on host use by a parasitic plant

Consumers do not always utilize all suitable hosts. Understanding why parasitic plants do not always parasitize potentially suitable hosts requires a better understanding of the constraints that limit host use by parasitic plants. In Texas salt marshes, the parasitic plant Cuscuta indecora rarely parasitizes three hosts that support vigorous growth in the greenhouse. We identified three constraints on host use by C. indecora. First, a mismatch between the phenology of C. indecora and some suitable hosts meant that these hosts were not abundant when C. indecora was growing most vigorously, and therefore were underutilized. Second, C. indecora preferentially parasitized tall plants versus short ones, causing relatively short species to be underutilized. Third, C. indecora overwinters in some perennial hosts but has to reinfect annual hosts each year, causing annuals and perennials that do not support overwintering to be underutilized. In combination, these constraints, which reflect the general lack of mobility of parasitic plants relative to herbivores, remove half of the potential host species from the actual diet of C. indecora, and therefore likely represent a major limitation on the success of this parasite. Similar constraints are likely to limit the realized host range of many parasitic plants and select for generalized diet preferences.

Biological control of Acacia cyclops in South Africa: The fundamental and realized host range of Dasineura dielsi (Diptera: Cecidomyiidae)

Acacia cyclops is one of the 13 species of Australian acacias (Mimosaceae) that have become naturalized in South Africa and are now declared invasive weeds. Dasineura dielsi, a multivoltine midge that induces galls on the ovary of its host, has been introduced for biological control of A. cyclops in South Africa where it now colonizes 100% of the mature trees and substantially reduces their reproductive capabilities. Although A. cyclops is the primary host, galls are sometimes induced by on several other Australian Acacia species, including Acacia longifolia, Acacia melanoxylon and Acacia saligna in South Africa. The mass of D. dielsi galls and adult midges and potential fecundity of the females were about the same on A. cyclops and on A. melanoxylon but were lower on A. longifolia and even lower on A. saligna. Cross-over experiments demonstrated that adults emerging from each host-plant species were capable of mating and producing viable progeny, confirming that these plants are within the fundamental host range of D. dielsi and that they have become incorporated into the realized host range of the midge in South Africa. In contrast to A. cyclops, these three alternate hosts each have relatively brief flowering periods which precede that of A. cyclops. This, in combination with the emergence pattern displayed by D. dielsi, seems to play a role in the extent to which the alternate hosts are utilized by D. dielsi. All indications are that D. dielsi will be of no significance on any species other than A. cyclops.

The ecological host range of an intentionally introduced herbivore: A comparison of predicted versus actual host use

An underlying assumption of weed biological control asserts that laboratory-based host specificity testing accurately predicts the realized host range of herbivorous arthropods. We tested this assumption by comparing predicted host specificity with the realized host range of the introduced weevil Oxyops vitiosa (Pascoe). Laboratory host specificity tests showed that the weevil can complete its development on only a small group of species in the Melaleuca genus, including the target host Melaleuca quinquenervia (Cav.) Blake. Three years after its release, adult weevils readily emigrated from the surrounding M. quinquenervia trees into replicated common gardens but only a small proportion occurred within the canopies of non- Melaleuca species. Adults remained within the canopies of Melaleuca congeners longer than non- Melaleuca and the former recruited 98% of all individuals dispersing from neighboring test plant species. Oviposition was predicted to occur on seven of the 19 species planted in the common gardens but was realized on only four exotic species. Consistent with the prerelease assessments, larvae were observed on four plant species but larval development was limited to Melaleuca congeners. These results lend support to the premise that risk assessments based on physiological host ranges, as characterized by laboratory testing, are conservative when compared to the realized ecological host ranges that occur under field conditions. We conclude that the ecological host range of O. vitiosa is highly restricted and there will be no colonization of species other than the target weed in the adventive range.

Laboratory and Realized Host Ranges of <I>Chaetorellia succinea</I> (Diptera: Tephritidae), an Unintentionally Introduced Natural Enemy of Yellow Starthistle

In 1999, we reported our discovery, in California and Oregon, of Chaetorellia succinea (Costa) (Diptera: Tephritidae) destroying the seeds of yellow starthistle, Centaurea solstitialis L., one of the worst weeds in the western United States. This fly, an unintentional introduction from Greece, dispersed rapidly throughout California and the northwest, and there is interest in using this adventive fly as a classical biological control agent for this weed. Because the host range of Ch. succinea has not been studied, this fly might pose a risk to other members of the thistle tribe Cardueae, especially the many thistle species native to California and other parts of the western United States. We determined the physiological host range of this fly in the laboratory by exposing it under no-choice conditions to 14 potential Cardueae hosts. Two introduced weed species and the native American basketflower (Centaurea americana Nuttall) were laboratory hosts. Under less restrictive choice test conditions, yellow starthistle was highly preferred, but there was a small amount of oviposition, and a few adult Ch. succinea emerged from all three of these plant species. Because Ch. succinea is now widespread throughout California, we collected flower heads from 24 potential host plant species at 111 sites to determine the realized host range in the field. These collections did not include American basketflower, which does not occur naturally in California. Ch. succinea emerged only from the other two known hosts: Ce. melitensis and Ce. sulfurea. Our results suggest that American basketflower growing in the southwestern United States may be at risk if Ch. succinea expands its range into that region.

Chemistry of Cirsium and Carduus: a role in ecological risk assessment for biological control of weeds?

Prediction of host plant range and ecological impact of exotic phytophagous insects, such as insects for classical biological control of weeds, represents a major challenge. Recently, the flowerhead weevil ( Rhinocyllus conicus Fröl.), introduced from Europe into North America to control exotic thistles ( Carduus spp.), has become invasive. It feeds heavily on some, but not all species of native North American thistles ( Cirsium spp.). We hypothesized that such non-target use among native plants could be better predicted by knowledge of characteristic chemical profiles of secondary compounds to supplement the results of host specificity testing. To evaluate this hypothesis, we reviewed the literature on the chemistry of Cirsium and Carduus thistles. We asked what compounds are known to be present, what is known about their biological activity, and whether such information on chemical profiles would have better predicted realized host range and ecological effects of R. conicus in North America. We found an extensive, but incomplete literature on the chemistry of true thistles. Two main patterns emerged. First, consistent chemical similarities and interesting differences occur among species of thistles. Second, variation occurs in biologically active groups of characteristic compounds, specifically flavonoids, sterols, alkaloids and phenolic acids, that are known to influence host plant acceptance, selection, and feeding by phytophagous insects. Surprisingly, sesquiterpene lactones, which are characterisitic in closely related Asteraceae, have not been extensively reported for Cirsium or Carduus. The minimal evidence on sesquiterpene lactones may reflect extraction methods vs. true absence. In summary, our review suggests further research on thistle chemistry in insect feeding is warranted. Also, since the exotic Canada thistle ( Cirsium arvense) is an invasive thistle of current concern in North America, such research on mechanisms underlying host range expansion by exotic insects would be useful.