Biological Control Of Invasive Plants Research Articles

Do thermal requirements of Dichrorampha odorata, a shoot-boring moth for the biological control of Chromolaena odorata, explain its failure to establish in South Africa?

Chromolaena odorata (L.) RM King and H Rob. (Asteraceae) has been subject to a biological control programme in South Africa for over three decades. A shoot-tip boring moth, Dichrorampha odorata Brown and Zachariades (Lepidoptera: Tortricidae), originating from Jamaica, was released as a biological control agent in 2013 but despite the release of substantial numbers of the insect, it has not established a permanent field population. Because climate incompatibility is a major constraint for classical biological control of invasive plants, and based on the differences in climate between Jamaica and South Africa and field observations at release sites, aspects of the thermal physiology of D. odorata were investigated to elucidate reasons for its failure to establish. Developmental time decreased with increasing temperatures ranging from 20 °C to 30 °C, with incomplete development for immature stages at 18 °C and 32 °C. The developmental threshold, t, was calculated as 8.45 °C with 872.4 degree-days required to complete development (K). A maximum of 6.5 generations per year was projected for D. odorata in South Africa, with the heavily infested eastern region of the country being the most eco-climatically suitable for establishment. The lower lethal temperature (LLT50) of larvae and adults was –4.5 and 1.8 °C, respectively. The upper lethal temperature (ULT50) for larvae was 39.6 °C whilst that of adults was 41.0 °C. Larvae thus had better cold tolerance compared to adults whereas adults had better heat tolerance compared to larvae. The critical thermal (CT) limits for adults were 3.4 ± 0.07 to 43.7 ± 0.12 °C. Acclimation at 20 °C for 7 days resulted in increased cold and heat tolerance with a CTmin and CTmax of 1.9 ± 0.06 and 44.4 ± 0.07 °C respectively, compared to the relative control, acclimated at 25 °C. Acclimation at 30 °C improved neither cold (CTmin: 5.9 ± 0.08 °C) nor heat tolerance (CTmax: 42.9 ± 0.10 °C). These results suggest that thermal requirements fall within field temperatures and are thus not the main constraining factor leading to poor establishment of D. odorata in South Africa.

Biological Control of Weeds: Is it Safe?

Invasive plants are non-native plant species that cause harm in their introduced range. Classical biological control of invasive plants is the use of natural enemies, imported insects and mites or pathogens, to control the target plants. This publication explains the strategies and rules in place to ensure that organisms released for the biological control of weeds are safe and effective.

Impacts of a biocontrol agent on invasive Ageratina adenophora in Southwest China: Friend or foe?

Classical biological control of invasive plants depends on the introduction of host-specific natural enemies. Many natural enemies have provided successful control, but the impacts of herbivorous insects on their hosts are highly variable and context dependent. Under some circumstances, overcompensation can even occur, with plant reproduction temporarily increasing rather than decreasing in response to herbivory. The stem-galling fly Procecidochares utilis has been widely introduced to help control Ageratina adenophora, a globally significant weed, but its impact has been inconsistent and in places ineffective. Its galling is known to stimulate production of side branches in plants under laboratory conditions. We examined the abundance and impact of P. utilis at four A. adenophora sites of Southwest China that were invaded at different times and support different densities of the insect. Stems with galls were more likely to be branched and produced more capitula. Furthermore, seed numbers in capitula from galled and un-galled stems were similar, and galling resulted in only slightly reduced seed weights. However, the increase in above-ground vegetative biomass associated with branching resulted in less efficient capitulum production relative to vegetative growth. Clearly Procecidochares utilis is not providing effective biocontrol of Ageratina adenophora in Southwest China so far. In the long term, galled plants may display reduced competitive ability and lower lifetime reproductive success, but the short term increases in seed production it generates suggest it may be favouring its host’s range expansion. The fly has been a successful biocontrol agent in dry area of Hawii, so understanding the factors that determine its effectiveness represent a major challenge for the future.

Climate change effects on biological control of invasive plants by insects.

Abstract Because many invasive plants are anticipated to benefit from climate change, the need for effective management strategies will become more pressing in the future. Biological control of invasive plants by insects has been an effective management strategy in many instances, but climate change may substantially alter such systems. Maximizing biological control efficacy with climate change will require predicting responses of the invasive plant in question, its biological control agents, and especially their corresponding interactions. However, because of the wide variety of species-specific plant and insect responses to increased atmospheric CO 2 concentrations, temperatures, extreme weather events, and other climate factors, climate change will likely result in highly variable, system- and geographically-specific effects. Even still, predicted future climates are generally anticipated across systems to alter geographic ranges of invasive plants (and correspondingly their biological agents), reduce host-plant quality, and hasten the phenology of both plants and insects. Some of these anticipated effects on biological control systems can be expected to hamper control efficacy. However, because some of these effects might also increase control efficacy, biological control can still be an effective and important management tool in the face of climate change. Although the variety of insect and plant responses to climate change reported in the literature do not currently allow for strong generalizations to be made as to the best biological control agents or strategies in novel climates, pertinent considerations for selecting effective biological control agents in the future are discussed.

First record of an apparently rare fig wasp feeding strategy: obligate seed predation

1. Fig trees require host‐specific agaonid fig wasps for pollination, but their figs also support numerous non‐pollinating fig wasps (NPFW) that gall fig tissues or develop as parasitoids.2. Ficus microcarpa L. is widely naturalised outside its native range and the most invasive fig tree species. Seed predators are widely used for the biological control of invasive plants, but no obligate seed predatory (as opposed to ovule or fig wall galling) NPFW have been recorded previously from any fig trees.3. Philotrypesis NPFW are usually regarded as parasitoids or ‘inquilines’ (parasitoids that also eat plant material) of pollinator fig wasps, but the present study provides evidence that Philotrypesis taiwanensis, a NPFW associated with F. microcarpa, is an obligate seed predator: (i) adults emerge from seeds of typical appearance, with a surrounding elaiosome; (ii) it shows no preference for figs occupied by fig wasp species, other than the pollinator; (iii) it only develops in figs that contain pollinated ovules, avoiding figs occupied by an agaonid that fails to pollinate; (iv) larvae are distributed in layers where seeds are concentrated and (v) it has a negative impact on seed but not pollinator offspring numbers.4. Philotrypesis is a large genus, and further species are likely to be seed predators.

Biocontrol attack increases pollen limitation under some circumstances in the invasive plant Centaurea solstitialis

Herbivore damage often deters pollinator visitation and many invasive plants in North America are pollinator-dependent. This has important implications for the biological control of invasive plants because it means that agents that deter pollinators may have a larger than expected impact on the plant. Yet interactions between pollinators and biocontrol agents are rarely evaluated. Centaurea solstitialis, one of the most problematic invasive species in California, is dependent on pollinators for reproduction. I factorially manipulated infection by a biocontrol pathogen and pollen supplementation to test for (1) pollen limitation in C. solstitialis, (2) whether infection increased pollen limitation, and (3) whether this varied across a soil moisture gradient. Plants growing on north-facing slopes where soil moisture was higher experienced mild pollen limitation in the absence of the pathogen and more pronounced pollen limitation when they were infected. Plants on drier south-facing slopes did not suffer from pollen limitation but instead appeared to suffer from resource limitation. Pathogen infection directly reduced seed set in C. solstitialis by 67-72%. On north-facing slopes, infection had an additional, indirect effect by increasing the degree of pollen limitation plants experienced. The trait that mediates this indirect pathogen-pollinator interaction is the number of inflorescences plants produced: infected plants made fewer inflorescences which led to greater pollen limitation. Although in the present study this outcome is dependent on abiotic factors that vary over small spatial scales, exploiting other invasive plants' dependence on pollinators by selecting agents that deter visitation may enhance agent impact.

Specificity of extrafloral nectar induction by herbivores differs among native and invasive populations of tallow tree.

Invasive plants can be released from specialist herbivores and encounter novel generalists in their introduced ranges, leading to variation in defence among native and invasive populations. However, few studies have examined how constitutive and induced indirect defences change during plant invasion, especially during the juvenile stage. Constitutive extrafloral nectar (EFN) production of native and invasive populations of juvenile tallow tree (Triadica sebifera) were compared, and leaf clipping, and damage by a native specialist (Noctuid) and two native generalist caterpillars (Noctuid and Limacodid) were used to examine inducible EFN production. Plants from introduced populations had more leaves producing constitutive EFN than did native populations, but the content of soluble solids of EFN did not differ. Herbivores induced EFN production more than simulated herbivory. The specialist (Noctuid) induced more EFN than either generalist for native populations. The content of soluble solids in EFN was higher (2·1 times), with the specialist vs. the generalists causing the stronger response for native populations, but the specialist response was always comparable with the generalist responses for invasive populations. These results suggest that constitutive and induced indirect defences are retained in juvenile plants of invasive populations even during plant establishment, perhaps due to generalist herbivory in the introduced range. However, responses specific to a specialist herbivore may be reduced in the introduced range where specialists are absent. This decreased defence may benefit specialist insects that are introduced for classical biological control of invasive plants.

Complex interactions among biocontrol agents, pollinators, and an invasive weed: a structural equation modeling approach

Herbivores, seed predators, and pollinators can exert strong impacts on their host plants. They can also affect the strength of each other's impact by modifying traits in their shared host, producing super- or sub-additive outcomes. This phenomenon is especially relevant to biological control of invasive plants because most invaders are attacked by multiple agents. Unfortunately, complex interactions among agents are rarely studied. We used structural equation modeling (SEM) to quantify the effect of two biocontrol agents and generalist pollinators on the invasive weed Centaurea solstitialis, and to identify and quantify the direct and indirect interaction pathways among them. The weevil Eustenopus villosus is both a bud herbivore and a predispersal seed predator; the fly Chaetorellia succinea is also a predispersal seed predator; Apis mellifera is the primary pollinator. We conducted this work at three sites spanning the longitudinal range of C. solstitialis in California (USA) from the coast to the Sierra Nevada Mountains. SEM revealed that bud herbivory had the largest total effect on the weed's fecundity. The direct effect of bud herbivory on final seed set was 2-4 times larger in magnitude than the direct effect of seed predation by both agents combined. SEM also revealed important indirect interactions; by reducing the number of inflorescences plants produced, bud herbivory indirectly reduced the plant's attractiveness to ovipositing seed predators. This indirect, positive pathway reduced bud herbivory's direct negative effect by 11-25%. In the same way, bud herbivory also reduced pollinator visitation, although the magnitude of this pathway was relatively small. E. villosus oviposition deterred C. succinea oviposition, which is unfortunate because C. succinea is the more voracious of the seed predators. Finally, C. succinea oviposition indirectly deterred pollinator visitation, thereby enhancing its net effect on the plant. This study demonstrates the powerful insights that can be gained from the SEM approach in understanding the multiple direct and indirect interactions among agents and pollinators and their effects on an invasive weed. Such an approach may improve our ability to manage weeds with biocontrol agents by identifying pathways that could be exploited by future agents and minimizing the possibility of interference with established agents.

The effectiveness of classical biological control of invasive plants

Summary Invasive alien plants have serious economic and ecological impacts, for example, by displacing native plants and invertebrates, and their management is often costly and ineffective in the long term. Classical biological control using specialized, coevolved natural enemies from the native region of the invader is often advocated as a preferred alternative to chemical and mechanical control, but there is a lack of quantitative assessment of control of the target species and subsequent establishment of native vegetation and invertebrates. Meta‐analyses were carried out combining the results of 61 published studies (2000–2011) that quantified the impact of classical biocontrol at the level of individual target plants, target populations or non‐target vegetation. Factors associated with the control programmes (invasive region, native region, plant growth form, target longevity, control agent guild, taxonomy and study duration) were analysed to identify patterns in control success. On average, biocontrol agents significantly reduced plant size (28 ± 4%), plant mass (37 ± 4%), flower and seed production (35 ± 13% and 42 ± 9%, respectively) and target plant density (56 ± 7%). Beetles in the Chrysomelidae and Curculionidae families were more effective at reducing plant size than other groups. Non‐target plant diversity significantly increased by 88 ± 31% at sites where biocontrol agents were released, but it was largely unclear whether the replacement plant species were native or invasive. Synthesis and applications. The number of studies that provide quantitative indications of the effectiveness of biocontrol and the response of non‐target taxa has increased over the past decade, but remains small compared to the total number of publications on the classical biocontrol of invasive plants. Nonetheless, this study demonstrates the positive impacts of classical biocontrol and the re‐establishment of native plants in a broad range of systems and establishes the value of classical biocontrol for the control of invasive alien plants. The Chrysomelidae and Curculionidae families were the most effective agents and we recommend these be prioritized in cases where potential agents of different taxa have also been identified. In addition, data on the recovery of native plant species and the invertebrate community remain sparse and it is recommended that future studies report the identity of plant species that replace target species as well as invertebrate community responses.

Biological control of invasive plants through collaboration between China and the United States of America: a perspective

For more than 100 years, classical biological control of invasive plants through screening, introducing and releasing of host-specific natural enemies from native regions has been regarded as one of the promising approaches to the management of invasive plants. Many invasive plants in the United States of America are native to China, and vice versa. China and the USA also share a number of invasive plant species, including water hyacinth (Eichhornia crassipes), alligator weed (Alternanthera philoxeroides) and cordgrass (Spartina spp.). Collaboration between the two countries on biological control benefits both the nations by reciprocal opportunities to research and exchange natural enemies, by exchanging information on common invasive species, and by providing training for students and professionals. Here we review the history of collaboration between China and the US on biological control of more than 20 invasive plants. Current collaborative projects associated with four plant species, Polygonum perfoliatum L., Trapa natans L. Pueraria montana (Lour.) Merr. var. lobata and Ailanthus altissima (Mill.) Swingle, are also covered. We prioritize 14 invasive plants as targets for future collaborative biological control based on information on their importance in introduced areas, natural enemy records, and their potential biocontrol risk to introduced ecosystems. They are: Ampelopsis brevipendunculata, Celastrus orbiculatus, Dioscorea oppositifolia, Euonymus alata, Euonymus fortunei, Ligustrum sinense, Melia azedarach, Paulownia tomentosa, Sapium sebiferum and Ulmus pumila for the US, as well as Spartina alterniflora, Ambrosia artemisiifolia, Ambrosia trifida and Solidago canadensis for China. In addition, we emphasize that we must very carefully consider any potential non-target effect when we intend to introduce and release new natural enemies. We anticipate that the high priority both countries have placed on control of invasive plants will stimulate increasing collaboration on biological control.