Peter G. Mason

Do new Access and Benefit Sharing procedures under the Convention on Biological Diversity threaten the future of biological control

Under the Convention on Biological Diversity (CBD) countries have sovereign rights over their genetic resources. Agreements governing the access to these resources and the sharing of the benefits arising from their use need to be established between involved parties [i.e. Access and Benefit Sharing (ABS)]. This also applies to species collected for potential use in biological control. Recent applications of CBD principles have already made it difficult or impossible to collect and export natural enemies for biological control research in several countries. If such an approach is widely applied it would impede this very successful and environmentally safe pest management method based on the use of biological diversity. The CBD is required to agree a comprehensive Access and Benefit Sharing process in 2010, in preparation for which the IOBC (International Organization for Biological Control of Noxious Animals and Plants) Global Commission on Biological Control and Access and Benefit Sharing has prepared this position paper. Here, we first describe the practice of biological control in relation to the principles of ABS, illustrated extensively by case studies and successes obtained with biological control. Next, we emphasise the very limited monetary benefits generated in biological control when compared to other fields of ABS such as the collection of germplasm for development of human drugs, chemical pesticides or crop cultivars. Subsequently, we inform the biological control community of good ABS practice and challenges, and we hope to make clear to the community involved in ABS under the CBD the special situation with regard to biological control. Finally, based on the non-commercial academic research model, we make recommendations which would facilitate the practice of collection and exchange of biological control agents, propose a workable framework to assist policy makers and biological control practitioners, and urge biological control leaders in each country to get involved in the discussions with their national ABS contact point to take their needs into consideration.

Benefits and risks of exotic biological control agents

The use of exotic (=alien) arthropods in classical and augmentative biological control programs has yielded huge economic and ecological benefits. Exotic species of arthropods have contributed to the suppression of key pests in agriculture and forestry or have aided in restoring natural systems affected by adventive species. However, adverse non-target effects of exotic biological control agents have been observed in a number of projects. Non-target effects range from very small effects, e.g. 2% parasitization on a non-target insect on a local level, to massive effects on a large scale. Until now, no consensus on how to judge the magnitude of non-target effects and whether these effects can be tolerated or are unacceptable has emerged. In this paper, we briefly review both the benefits of biological control as well as the associated risks including to human and animal health, plant health and particularly the environment. We also make an attempt at identifying the major challenges for assessing risks and for balancing benefits and risks. There is general agreement that sound risk assessment procedures should precede the release of exotic invertebrate biological control agents and a recent shift—especially for arthropod biological control—from introductions done without meaningful risk assessment studies to projects conducting thorough host range testing can be observed. However, overly stringent regulations that would preclude promising agents from being developed must be avoided.

DNA barcoding and the taxonomy of Microgastrinae wasps (Hymenoptera, Braconidae): Impacts after 8 years and nearly 20 000 sequences

Microgastrine wasps are among the most species‐rich and numerous parasitoids of caterpillars (Lepidoptera). They are often host‐specific and thus are extensively used in biological control efforts and figure prominently in trophic webs. However, their extraordinary diversity coupled with the occurrence of many cryptic species produces a significant taxonomic impediment. We present and release the results of 8 years (2004–2011) of DNA barcoding microgastrine wasps. Currently they are the best represented group of parasitoid Hymenoptera in the Barcode of Life Data System (BOLD), a massive barcode storage and analysis data management site for the International Barcoding of Life (iBOL) program. There are records from more than 20 000 specimens from 75 countries, including 50 genera (90% of the known total) and more than 1700 species (as indicated by Barcode Index Numbers and 2% MOTU). We briefly discuss the importance of this DNA data set and its collateral information for future research in: (1) discovery of cryptic species and description of new taxa; (2) estimating species numbers in biodiversity inventories; (3) clarification of generic boundaries; (4) biological control programmes; (5) molecular studies of host‐parasitoid biology and ecology; (6) evaluation of shifts in species distribution and phenology; and (7) fostering collaboration at national, regional and world levels. The integration of DNA barcoding with traditional morphology‐based taxonomy, host records, and other data has substantially improved the accuracy of microgastrine wasp identifications and will significantly accelerate further studies on this group of parasitoids.

Stable Bacillus thuringiensis (Bt) toxin content in interspecific F1 and backcross populations of wild Brassica rapa after Bt gene transfer

Stable expression of a transgene may lead to increased fitness for wild plants after acquiring the transgene via crop–weed hybridization. Here, we investigate the stability of Bt toxin content in wild Brassica rapa acquiring the Bt gene from Bt Brassica napus. The Bt toxin content in nine Bt‐expressing B. napus lines was 0.80–1.70 µg/g leaf tissue throughout the growing season. These nine lines were crossed with three accessions of wild B. rapa and the Bt gene was successfully transferred to interspecific hybrids (F1) and successive backcross generations (BC1 to BC4). The Bt toxin level in F1 and BC progenies containing the Bt gene remained at 0.90–3.10 µg/g leaf tissue. This study indicates that the Bt gene can persist and be stably expressed in wild B. rapa.

Key Pests and Parasitoids of Oilseed Rape or Canola in North America and the Importance of Parasitoids in Integrated Management

The development of ‘double low oilseed rape’ or canola as a unique crop in the mid 1970s resulted in rapid expansion in the area devoted to its production in North America, especially in the western Canadian provinces of Alberta, Saskatchewan and Manitoba. The crop was soon subjected to attack by a number of insect herbivores, sometimes responsible for severe economic losses. The key pests of the crop comprise the flea beetles Phyllotreta cruciferae (Goeze) and Phyllotreta striolata (Fabricius), several root maggot species especially Delia radicum (L.), cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham), bertha armyworm, Mamestra configurata Walker, several species of lygus bugs, Lygus spp., and the diamondback moth, Plutella xylostella (L.). Most pest species are alien, introduced from Europe or Asia, and as a consequence they have been the subject of importation and release of non-indigenous parasitoids in attempts to achieve classical biological control. To date, such attempts have met with limited success, but nevertheless substantial progress has been made in certain situations. This chapter reviews the life history and crop damage caused by each of the key pest species, and the parasitoid fauna associated with each. Attempts at classical or augmentative biological control of the pests are documented, and a case history is provided for one invasive species, the cabbage seedpod weevil. Challenges to biological control with parasitoids are identified for North American cropping systems.

Importance of long-term research in classical biological control: an analytical review of a release against the cabbage seedpod weevil in North America

Abstract:  Cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (Col., Curculionidae), is an invasive alien pest that is spreading in North America. To aid with planning for introductions of European parasitoids in North America, we examined the status of the only classical biological control release against this pest in North America, which in 1949 introduced Mesopolobus morys, Stenomalina gracilis and Trichomalus perfectus (Hym., Pteromalidae). Weevils and parasitoids were reared in 2005 from mass collections of seedpods of Brassica napus, Brassica rapa and Raphanus raphanistrum (Brassicaceae) from 18 sites in the Fraser Valley, near Vancouver, British Columbia, Canada. Of the three European parasitoid species that were originally released, only S. gracilis was found. The predominant hymenopterous parasitoid species were Trichomalus lucidus, S. gracilis, Mesopolobus moryoides (Pteromalidae), Necremnus tidius (Eulophidae) and Eupelmus vesicularis (Eupelmidae). These constituted over 97% of the parasitoids reared, although overall parasitism was low. Only M. moryoides is clearly North American in distribution; other than S. gracilis, the remaining species were either accidentally introduced or are Holarctic in distribution. Based on these results, re‐releases of M. morys and T. perfectus in North America should be considered as part of a classical biological control programme. However, redistribution of S. gracilis is not recommended at present because of potential conflicts with biological control programmes against weeds. Ongoing re‐examination of classical biological control programmes can further our understanding of failure of release programmes, particularly when re‐examination can be made in the light of improved taxonomy and systematics of the target and agent species.

Controlling Lygus plant bugs (Heteroptera: Miridae) with European Peristenus relictus (Hymenoptera: Braconidae) in Canada - risky or not?

The European Peristenus relictus Loan (syn. P. stygicus) has been considered for biological control of Lygus plant bugs native to Canada. Laboratory and field studies were conducted in the area of origin to evaluate the host specificity of P. relictus. Laboratory choice and no-choice tests demonstrated that P. relictus attacked all non-target species offered (fundamental host range). However, closely related non-target mirids (tribe Mirini) were generally well accepted by P. relictus, while hosts from the tribe Stenodemini were less frequently attacked and less suitable for parasitoid development. To validate the laboratory results, a thorough examination of the parasitoid complex of common mirids in Europe was conducted to determine which non-target species may serve as alternative hosts for P. relictus in a natural situation (ecological host range). When comparing both approaches, the fundamental host range of P. relictus matched its ecological host range. In addition to three Lygus species, the ecological host range of P. relictus in the area of Schleswig-Holstein, northern Germany, contains at least 16 non-target species, including hosts belonging to the subfamilies Mirinae, Phylinae and Bryocorinae. A broad ecological and fundamental host range suggests that P. relictus is a generalist; however, P. relictus was not the primary contributor to parasitism of most non-target hosts studied. Although P. relictus is assumed to be of minor importance for regulating non-target populations in the area of investigation, the results of the present study indicate that P. relictus has the potential to use non-target host populations for reproduction.

Interaction of a solitary larval endoparasitoid, Microplitis mediator, with its host, Mamestra brassicae: host acceptance and host suitability

Abstract:  Microplitis mediator (Haliday) (Hym., Braconidae) is an important parasitoid of early instar larvae of the European cabbage moth, Mamestra brassicae L. (Lep., Noctuidae). In the laboratory, we examined attack responses of female M. mediator to the first three larval instars of M. brassicae. Females were presented with M. brassicae larvae either one individual at a time in a no‐choice experiment, or three individuals, one from each instar, simultaneously in a choice experiment. Whether or not there was choice, naïve female parasitoids attacked a high proportion of larvae and did not discriminate among instars. In the no‐choice experiment, attacked larvae were reared, and parasitoid cocoons were produced from about 76% of larvae attacked as first and second instars, but from only 19% of larvae attacked as third instars. Dissections of attacked larvae from the choice experiment showed that about 79% of attacks on first and second instars resulted in oviposition compared with only 49% for third instars. When given choice, frequency and number of attacks on first instar larvae increased with increasing parasitoid experience. Our results suggest that first and second instar larvae of M. brassicae are suitable hosts for M. mediator, but that third instar larvae are suboptimal both because oviposition attempts were frequently unsuccessful and because immature parasitoids failed to complete development. Nevertheless, naïve attacking parasitoids exhibited minimal discrimination among instars, although experienced parasitoids most frequently attacked first instar larvae. The host selection behaviour of M. mediator is discussed in the context of optimal foraging theory and implications for biological control.

A review of the species of Mesopolobus (Chalcidoidea: Pteromalidae) associated with Ceutorhynchus (Coleoptera: Curculionidae) host-species of European origin

Four species of Mesopolobus Westwood were reared as parasitoids of Ceutorhynchinae hosts in Europe during surveys in 2000-2004. An illustrated key is given to differentiate the four species, M. gemellus Baur & Muller sp. n., M. incultus (Walker), M. morys (Walker) and M. trasullus (Walker), plus M. moryoides Gibson, a parasitoid of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham), in North America. Pteromalus clavicornis Walker is recognized as a junior synonym of M. incultus syn. n., and Pteromalus berecynthos Walker (also a junior synonym of M. incultus) is considered a correct original spelling. For Disema pallipes Förster (a junior synonym of Mesopolobus morys), a lectotype is designated. Mesopolobus morys is for the first time accurately associated with the seed weevil Ceutorhynchus turbatus (Schultze), a potential agent for classical biological control, of hoary cress, Lepidium draba L. (Brassicaceae), in North America. Mesopolobus gemellus is associated with another seed weevil, Ceutorhynchus typhae (=C. floralis) (Herbst), in pods of shepherds purse, Capsella bursa-pastoris (L.) Medik. (Brassicaceae). Implications of the host-parasitoid associations are discussed relative to the introduction of species to North America for classical biological control of the cabbage seedpod weevil.

Responses of Chalcidoidea (Hymenoptera) parasitoids to invasion of the cabbage seedpod weevil (Coleoptera: Curculionidae) in western Canada

Invasion of the European weevil, Ceutorhynchus obstrictus (Marsham), was investigated through surveys of its range and population densities in Alberta and Saskatchewan from 2001 to 2005. After it was first reported in southern Alberta, C. obstrictus rapidly expanded its range and abundance. Our more recent surveys indicate that its northward expansion has slowed, but that it has continued to extend its range eastward to southcentral Saskatchewan. The distribution and abundance of parasitoids of C. obstrictus in Alberta and Saskatchewan were investigated from 2003 to 2005 by mass rearing canola pods infested with C. obstrictus larvae. Although weevil populations were not parasitized for several years immediately following its introduction to southern Alberta, a surprisingly diverse assemblage of Chalcidoidea parasitoids, comprising 12 species from four families, were recently reared from weevil-infested canola siliques in Alberta and Saskatchewan. The Chalcidoidea fauna of C. obstrictus include species with both Nearctic and Holarctic distributions, with some species having restricted host ranges and others that are more niche than taxon-specific. These Chalcidoidea species appear to have expanded their host ranges to parasitize C. obstrictus in the region. Most parasitism is attributable to Trichomalus lucidus (Walker), Chlorocytus sp., and Pteromalus sp. (Pteromalidae), and Necremnus tidius (Walker) (Eulophidae). Parasitism levels varied considerably over the three years of this study. From 2003 to 2005 increases in parasitism occurred among all four of the species dominating the parasitoid fauna of C. obstrictus, but greater increases were observed for Chlorocytus sp. and Pteromalus sp. than for T. lucidus. Parasitoid species have sometimes caused substantial levels of host mortality, although current levels are usually less than 15% for all species combined and so are not sufficient to control weevil populations. Implementing a classical biological control program for C. obstrictus by reconstructing its European natural enemy complex is being considered, but it is still uncertain whether parasitism levels by native Chalcidoidea will increase over time since considerable year-to-year variation has been found. Parasitism levels of C. obstrictus should therefore continue to be monitored to assess whether a classical biological control program should be implemented.