Robert S. Bourchier

The influence of matrix habitat on Aphthona flea beetle immigration to leafy spurge patches

Variation in movement ability by insects among different non-habitat (matrix) types may have important implications for both metapopulation dynamics and weed biocontrol practices. We used a mark-recapture experiment to explore the effects of two different matrix habitats (grass vs shrub) on the ability of two species of Aphthona (Chrysomelidae: Coleoptera) flea beetle to immigrate to patches of the invasive weed, leafy spurge. Using generalized linear models, we compared effects of the matrix habitat types, species and sex on observed immigration probabilities. Our analyses demonstrated that one species (A. nigriscutis) had a much higher immigration probability when moving through a grass-dominated matrix than a shrub-dominated matrix whereas immigration probabilities for the second species (A. lacertosa) were similar in both matrix habitats but significantly lower overall than for A. nigriscutis. Furthermore, A. nigriscutis females were more likely to immigrate to spurge patches embedded in a grass matrix than in shrub, whereas the opposite occurred for males. Our results suggest that metapopulation dynamics may be strongly affected by the type(s) of matrix habitat present on a landscape. These effects also suggest that release strategies for weed biocontrol should be tailored according to the structure of the landscape into which releases are planned. In addition, even closely related species can have significantly different movement abilities which will also affect release strategies.

Polyploidy in Phenotypic Space and Invasion Context: A Morphometric Study of Centaurea stoebe s.l.

The taxonomy of the Centaurea stoebe complex is controversial. Diploid and tetraploid plants occur in its native European range, but to date only tetraploids have been recorded from its introduced range in North America. We examined morphological differentiation of C. stoebe using multivariate and univariate approaches to clarify the taxonomic status of the known cytotypes. We measured more than 40 morphological traits on plants originating from 78 populations, grown from seed under uniform glasshouse conditions. The ploidy of almost 300 plants from 2 native and 20 introduced populations from Canada was assessed to test for the absence of diploids from North America. Finally, we explored whether postintroduction processes have resulted in phenotypic changes in introduced plants which may have contributed to the invasion success of C. stoebe. Morphometric analyses showed a clear separation of 2x and 4x plants and thus supported recognition of both cytotypes as separate taxa. Differences in the life cycle, the number of florets, the shape of capitula, and the shape of young rosette leaves were the best discriminant characters. Only minor differences were found between native and introduced tetraploids. All plants from the introduced range except for one hexaploid were found to be tetraploid. Rare diploids from Canada were identified as Centaurea diffusa or Centaurea psamogenna.

Interspecific interactions between the gall-fly Urophora affinis Frfld. (Diptera: Tephritidae) and the weevil Larinus minutus Gyll. (Coleoptera: Curculionidae), two biological control agents released against spotted knapweed, Centaurea stobe L. ssp. micranthos

Abstract Interspecific competition has been suggested as an explanation for the failure of some insects as biological control agents for weeds. Enclosure and exclusion cages were used, in southern British Columbia, Canada to evaluate the importance of interspecific competition between a seedhead weevil, Larinus minutus, and a gall-inducing fly, Urophora affinis, two biocontrol agents released against spotted knapweed in North America. At the seedhead scale, U. affinis, which is an inferior biological control agent based on knapweed seed mortality, was the superior competitor. Larinus minutus attack rates were significantly lower in the presence of U. affinis compared to release treatments where L. minutus was attacking alone. Reduced L. minutus attack rates were apparent in seed heads expected to contain both species, assuming insect distributions were random, but instead only contained U. affinis. L. minutus did not significantly affect U. affinis density. Although overall attack rates on knapweed seedheads were higher when both species were together at a site, the consequence of the antagonistic interaction is that overall seed destruction was not as high as it could have been if the weevil were attacking on its own. These results support minimizing the number of biocontrol agents released that use similar resources on the target weed, to avoid negative interactions between control agents and potential reductions in biocontrol efficacy resulting from competitive exclusion.

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

Abstract In 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 agents 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 Groups evaluation process of biocontrol agents; (2) recognition of the primacy of realized host range data for potential agents that considers the insects 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.

Early establishment and dispersal of the weevil, Mogulones cruciger (Coleoptera: Curculionidae) for biological control of houndstongue (Cynoglossum officinale) in British Columbia, Canada

First released in Canada in 1997 to control the invasive rangeland weed, houndstongue (Cynoglossum officinale), the European root weevil, Mogulones cruciger, is showing early potential as a successful biocontrol agent. Out of 22 experimental releases in southeastern British Columbia, Canada, 100% established, regardless of initial release size (range 100–400). These founding populations persisted beyond 2 years, and quickly dispersed through a treed and variable landscape to colonise new houndstongue patches surrounding the original release patches. Within 3 years, the weevil had moved 1.42 km. Both initial, within-patch and later, between-patch dispersal followed a similar pattern, with M. cruciger adults and/or their feeding/oviposition damage being more concentrated nearest the original points of release and declining with distance. Within-patch, this pattern of distribution and also the rate of spread of weevils were similar regardless of initial release size; suggesting density-independence in dispersal behaviour at this scale. Closer investigation of the sequence of invasion at a larger spatial scale suggested that the weevil arrived at, colonized and increased in number on the closest host patches first. Three years post release, both distance from release patch and the number of M. cruciger released, were significant predictors of the amount of feeding/oviposition damage, and presumably weevil population size, within newly colonized houndstongue patches surrounding the original releases. These data, and the finding that more weevils were retrieved from patches where higher numbers of weevils were released in the previous year (i.e., 300 or 400 vs 100 or 200), suggest that release number and placement of releases within a landscape can be manipulated for effective houndstongue control.

Distribution and Potential Spread of Japanese Knotweed (Polygonum cuspidatum) in Canada Relative to Climatic Thresholds

Abstract Japanese knotweed (JK) is one of the most aggressive invasive plants known in the U.K., where its biology has been well-studied. It was introduced into Canada around 1900, but only recently has it become a serious concern in the province of British Columbia (BC). Climatic conditions, including annual degree days and mean-annual minimum temperatures at knotweed sites in British Columbia were modeled in BioSIM, using weather normals and long-term daily weather data, and compared to published thresholds (degree day  =  2,505 DD, minimum temperature  =  −30.2 C, base temperature 0 C ). The degree-day threshold was more limiting to JK in British Columbia than mean-minimum temperature (12.3% of province habitat was suitable for JK based on degree days compared with 26% for mean-minimum temperature). A new annual-precipitation threshold of 735 mm/year based on 95% of known knotweed sites in BC was identified. The best-fit logistic regression model included degree days and annual precipitation and predicted knotweed presence/absence with over 97% efficiency. Existing knotweed sites occupy just over half of the suitable habitat in BC, indicating there are still significant areas to be invaded. The limiting threshold for knotweed was reversed in Southern Ontario with between 35 to 53% of the habitat suitable based on minimum temperatures, whereas degree-day accumulations and annual precipitation were not limiting. Warmer temperatures from 2000 to 2008 resulted in an increase to 53% of the habitat in Southern Ontario being suitable for knotweed, compared to 35% when 1971 to 2000 weather normals were used. Different climatic thresholds among provinces might result in selection for different invasive knotweed genotypes. This could influence the success of biological control agents because of differential host suitability of knotweed genotypes. Habitat suitability maps generated will enable better targeting of knotweed surveys based on the risk of knotweed establishment. Nomenclature: Japanese knotweed, Polygonum cuspidatum Sieb. et Zucc. [syn. Fallopia japonica (Houtt.) Ronse Decraene var. japonica]; Giant knotweed, Polygonum sachalinense F. Schmidt ex Maxim.[syn. Fallopia sachalinensis (F. Schmidt) Ronse Decraene]; Bohemian knotweed, Polygonum ×bohemicum (J. Chrtek & Chrtková) Zika & Jacobson [cuspidatum × sachalinense] [syn. Fallopia × bohemica (Chrtek and Chrtková) J. P. Bailey.].

Effect of matrix habitat on the spread of flea beetle introductions for biological control of leafy spurge

Matrix habitats are known to influence the movement patterns of a variety of species but it is less well known whether these effects have strong implications for spatial population dynamics, including the spread of biological introductions. Using a spatially explicit simulation model parameterized with empirical data, we examine how grass and shrub matrix habitats, each offering different resistance to dispersal, influence the spread and impact of a biocontrol agent, Aphthona lacertosa, on the invasive weed, leafy spurge. Model predictions indicate that differential responses to matrix habitat have little effect on the agent’s spread over the study landscape and this is supported by statistical models fit to observed A. lacertosa incidence on the same landscape. Subsequent experimentation with the simulation model suggested that A. lacertosa colonization rates were largely unaffected by increases in amount of the more restrictive shrub matrix. However, simulations of an hypothetical species with greater overall dispersal ability but reduced dispersal rate through shrub matrix showed that colonization rates were noticeably reduced when the percentage of shrub matrix on the landscape approached 50%. Combined these results suggest that some tailoring of release strategies may be required to accommodate the unique dispersal capabilities of different biocontrol agents on particular release landscapes, but for A. lacertosa there appears to be little effect of matrix habitat structure on rates of spread.

Is two company or a crowd: How does conspecific density affect the small-scale dispersal of a weed biocontrol agent?

To predict the growth and spread of an insect population introduced for the biological control of weeds, one must first understand the factors affecting the movement of individuals in the population. The purpose of this study was to determine how the dispersal rate of Aphthona lacertosa (Rosenhauer) (Chrysomelidae) was affected by conspecific density and by the characteristics of leafy spurge (Euphorbia esula L.: Euphorbiaceae) in patches where these beetles feed. In 2002 in Manitoba and in 2003 in Alberta, Canada, between 200 and 2500 insects were released in small patches (<10 m2) of spurge. The number and location of beetles within patches was monitored over subsequent days. In 1 m2 plots within patches, spurge ramet density, the proportions of vegetative and reproductive ramets, and ramet height were measured. In both years, beetle movement within patches and emigration from patches, was not affected by conspecific density. In Manitoba in 2002, beetles aggregated non-randomly on either vegetative or reproductive ramets within plots, but plot characteristics were not related to the formation of aggregations. In Alberta in 2003, plots in which beetles aggregated had significantly higher spurge density but did not differ in either the proportion of vegetative ramets or in the amount of non-spurge vegetation. These results suggest that density-dependent dispersal does not limit the populations ability to reach densities up to 2500 beetles/m2.

Phylogeny and Genetic Diversity of Flea Beetles (Aphthona sp.) Introduced to North America as Biological Control Agents for Leafy Spurge

ABSTRACT A molecular phylogeny is presented for the five main species of Aphthona flea beetles that were introduced to North America in conjunction with the leafy spurge (Euphorbia esula L.) biological control program. The mitochondrial genome was examined using polymerase chain reaction-restriction fragment length polymorphism (RFLP) of a 9,000-bp segment and nucleotide sequencing of a 575-bp piece of cox1-cox2. A neighbor—joining tree of the RFLP data, along with neighbor-joining and maximum parsimony trees of the sequence alignments, all had the same major branching pattern. Each of the recognized species was a well defined clade. Three within species subbranches had very limited mitochondrial DNA diversity. One was a Wolbachia-infected lineage of A. nigriscutis most likely generated by a Wolbachia sweep where the spreading Wolbachia infection brought along the infected mitochondrial haplotype. Two of three subclades of A. lacertosa also had very little genetic diversity. One of these subclades also displayed a divergence from the other two that was analogous to the divergence observed between some of the other species pairs, suggesting it may be a cryptic species. Its distribution was restricted to Canada. The other genetically depauperate A. lacertosa line was the only lineage recovered in the United States. The geographically restricted nature of some of the genetic lines could be exploited to possibly improve biological control in some habitats through redistribution to other locations. It is not obvious that either Wolbachia infection or a narrow genetic base has had any detrimental effect on biological control.