Host specificity ofDisonycha argentinensis [Col.: Chrysomelidae], an agent for the biological control ofalternanthera philoxeroides (Alligator weed) in Australia

The biology and host specificity of a South American flea beetle,Disonycha argentinensis Jacoby was investigated in quarantine facilities in Australia. Damage from feeding by larvae and adults ofD. argentinensis onAlternanthera philoxeroides (Mart.) Griseb (alligator weed) indicated that it might be a useful biological control agent for the plant when growing in terrestrial situations. Complete development of the immature stages took place only onAlternanthera philoxeroides. Adults produced a few feeding scars onBeta vulgaris L. (beetroot) andAmaranthus spinosus L. (spiny amaranth) but failed to oviposit. Host specificity studies indicated that establishment of this beetle in Australia would be without significant risk to nontarget plant species.D. argentinensis was first released in Australia in 1980.     

Flooding compromises compensatory capacity of an invasive plant: implications for biological control

Plant compensatory growth is proposed to be insidious to biological control and known to vary under different environmental conditions. However, the effects of microsite conditions on compensation capacity and its indirect impacts on biological control of plant invaders have received little attention. Alligator weed, Alternanthera phioxeroides, is an invasive plant worldwide, growing in both aquatic and terrestrial habitats that are often affected by flooding. Biological control insects have been successful in suppressing the plant in many aquatic habitats but have failed in terrestrial habitats. To evaluate the impact of flooding on compensation capacity, we conducted common garden and greenhouse experiments in which plants were grown under different moisture conditions (aquatic versus terrestrial). Our results show that plants were able to fully recover from continued herbivory in the terrestrial habitat, but failed in the aquatic habitat, indicating a flooding-regulated plant compensatory capacity. Also, the grazed plants increased below-ground growth and reproductive root bud formation in the terrestrial habitat, but there was no such difference in the aquatic habitat. Our findings suggest that the differing plant compensatory capacity, affected by flooding, may explain the different biological control efficacy of alligator weed in aquatic and terrestrial habitats. Understanding mechanisms in plant invader compensation in different microsite conditions is important for improving management efficiency.     

Effects of simulated herbivory and resource availability on the invasive plant, Alternanthera philoxeroides in different habitats

In biological control programs, the insect natural enemy’s ability to suppress the plant invader may be affected by abiotic factors, such as resource availability, that can influence plant growth and reproduction. Understanding plant tolerance to herbivory under different environmental conditions will help to improve biocontrol efficacy. The invasive alligator weed (Alternanthera philoxeroides) has been successfully controlled by natural enemies in many aquatic habitats but not in terrestrial environments worldwide. This study examined the effects of different levels of simulated leaf herbivory on the growth of alligator weed at two levels of fertilization and three levels of soil moisture (aquatic, semi-aquatic, and terrestrial habitats). Increasing levels of simulated (manual) defoliation generally caused decreases in total biomass in all habitats. However, the plant appeared to respond differently to high levels of herbivory in the three habitats. Terrestrial plants showed the highest below–above ground mass ratio (R/S), indicating the plant is more tolerant to herbivory in terrestrial habitats than in aquatic habitats. The unfertilized treatment exhibited greater tolerance than the fertilized treatment in the terrestrial habitat at the first stage of this experiment (day 15), but fertilizer appears not to have influenced tolerance at the middle and last stages of the experiment. No such difference was found in semi-aquatic and aquatic habitats. These findings suggest that plant tolerance is affected by habitats and soil nutrients and this relationship could influence the biological control outcome. Plant compensatory response to herbivory under different environmental conditions should, therefore, be carefully considered when planning to use biological control in management programs against invasive plants.     

Control and spread of Alligator Weed Alternanthera philoxeroides (Mart.) Griseb., in Australia: lessons for other regions

Biological control of alligator weed Alternanthera philoxeroides (Mart.) Griseb. has been successful in limiting growth in water in areas with mild or warm winters, but not on land. Until recently, herbicides have had very limited short term and no long term effectiveness. Several herbicides that now provide better control include: glyphosate over water, and metsulfuron and dichlobenil on land and in shallow water. The latter two are limited by lack of selectivity, contamination of water, and cost. Mechanical or manual control has provided local eradication of the weed at a few locations where infestations were small. Alligator weed is still spreading with new outbreaks on New South Wales, Australia (NSW) coastal beach areas and coastal river systems, and on inland waterbodies. Its use as a cultivated vegetable by some ethnic communities has resulted in many new locations in all eastern Australia states: Queensland to Tasmania. It is predicted that it will spread throughout much of coastal and inland southern Australia. The difficulties with management of this weed indicate that every effort should be made to prevent further invasion of wetlands and, in particular, its introduction to Africa, where it is predicted that all wetlands could support destructive levels of alligator weed growth.     

Biological control of water lettuce,Pistia stratiotes [Araceae] byNeohydronomus affinis [Coleoptera: Curculionidae]

Neohydronomus affinis was imported into Australia as a potential biological control agent for the floating aquatic weedPistia stratiotes. Quarantine studies demonstrated that this weevil was host specific toP. stratiotes and it was released into the field in 1982. Subsequently it was released as a control agent for this weed in several other countries. In AustraliaN. affinis reduced infestations ofP. stratiotes by 40% or more within 12–18 months. The results are discussed in relation to its effectiveness in other parts of the world. We concluded thatN. affinis effectively controlsP. stratiotes in tropical regions (to lat. 22°), but in cooler regions its effectiveness fluctuates with seasonal conditions.      

Field assessment of a frond-feeding weevil, a successful biological control agent of red waterfern, Azolla filiculoides, in southern Africa

Azolla filiculoides (red waterfern) is a small, floating fern native to South America, that has invaded aquatic habitats, predominantly water resevoirs in southern Africa. A frond-feeding weevil, Stenopelmus rufinasus Gyllenhal (Coleoptera: Curculionidae), was imported from Florida, USA, and released as a biological control agent against this weed in South Africa at the end of 1997. To date, 24,700 weevils have been released, which has resulted in local extinction of red waterfern at 81% of the 112 release sites. The weevil has not failed to control a single site. Several sites were, however, lost due to flooding or drainage of dams. The surface area of weed controlled totalled 203.5 ha. On average, A. filiculoides was controlled in infested sites in 6.9 (±4.3) months. The weed recolonized at 22 of the sites (through either spore germination or dispersal by waterfowl), but the weevils subsequently spread to all of these sites and successfully caused local extinction of the weed at 18 of the sites. Five years after the release of the weevil, the weed no longer poses a threat to aquatic systems in southern Africa. In comparison to other biological control programs of aquatic weeds, the program against A. filiculoides in southern Africa ranks among the most successful cases anywhere in the world.     

Phenotypic plasticity rather than locally adapted ecotypes allows the invasive alligator weed to colonize a wide range of habitats

Both phenotypic plasticity and locally adapted ecotypes may contribute to the success of invasive species in a wide range of habitats. Here, we conducted common garden experiments and molecular marker analysis to test the two alternative hypotheses in invasive alligator weed (Alternanthera philoxeroides), which colonizes both aquatic and terrestrial habitats. Ninety individuals from three pairs of aquatic versus terrestrial populations across southern China were analyzed, using inter simple sequence repeat (ISSR) marker, to examine population differentiation in neutral loci. Two common gardens simulating aquatic and terrestrial habitats were set up to examine population differentiation in quantitative traits. We found no evidence of population differentiation in both neutral loci and quantitative traits. Most individuals shared the same ISSR genotype. Meanwhile, plants from different habitats showed similar reaction norms across the two common gardens. In particular, plants allocated much more biomass to the belowground roots in the terrestrial environment, where alligator weed may lose part or all of the aboveground shoots because of periodical or accidental disturbances, than those in the aquatic environment. The combined evidence from molecular marker analysis and common garden experiments support the plasticity hypothesis rather than the ecotype hypothesis in explaining the adaptation of alligator weed in a wide range of habitats.     

Risk to non-target plants from Charidotis auroguttata (Chrysomelidae: Coleoptera), a potential biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae) in Australia

The potential of the leaf beetle Charidotis auroguttata as a biocontrol agent for cat’s claw creeper Macfadyena unguis-cati (Bignoniaceae), an environmental weed in Australia, and risk to non-target plants was evaluated under quarantine conditions. In no-choice tests, C. auroguttata adults and larvae fed on many plant species across different families, but egg to adult development occurred only on the target weed. However, when neonate larvae from the target weed were transferred onto Myoporum boninense australe (Myoporaceae), a non-target native plant, 11.7% completed development, as compared to 95% of larvae that completed development on the target weed. Larval development on this non-target species also took twice as long as on the target weed. No larvae completed development on other test plants. In choice tests, leaf area consumption by adults and larvae was significantly more on the target weed than on other plants, and oviposition occurred only on the target weed. In the no-choice demography trials, adults laid eggs from the second week after emergence on the target weed, with an average of 0.286 eggs/female/day, resulting in an 18-fold increase in the adult population over 16 weeks. On My. boninense australe adult survival remained high, but oviposition commenced only from the 10th week after emergence with an average of 0.023 eggs/female/day, and none of the eggs developed into adults. In the choice demography trials, oviposition on the target weed was evident from the fourth week onwards, while on the non-target plant oviposition commenced only from the 14th week. Only 10% of total adults and 11.3% of total eggs were found on the non-target plant, and none of these eggs developed into adults. Although the biocontrol agent can ‘spill-over’ from the target weed to the non-target native plant and cause adult feeding damage, the non-target plant could not sustain a viable insect population on its own. This agent was not approved for field release in Australia due to perceived risk to non-target species.     

Natural Enemies ofTrapaspp. in Northeast Asia and Europe

Trapa natansis an aquatic plant native to the Old World. In the late 1880s this plant was found to be naturalized in the eastern United States where it has become an important weed. To evaluate the potential of biological control to reduce this weed, surveys for natural enemies ofTrapaspp. were made in Northeast Asia and Europe. In addition, populations ofTrapa japonicain South Korea were monitored for one season to evaluate the occurrence and impact of insect natural enemies. Among the insects found, the leaf beetleGalerucella birmanicawas the most common and damaging species in Asia. It can cause complete defoliation of whole populations of plants. Nymphuline pyralid moths were also common and at times damaging. Both the beetle and the moths feed and develop on unrelated plants and so have no potential as biological control agents ofT. natansin North America. TwoNanophyesweevils were found in Asia which feed in petiole floats of the leaves. They are thought to be specific toTrapabut were not observed to be damaging. Low level populations of polyphagous Homoptera were common. Chironomid midges were also frequently associated with the plants, but for the most part were filter feeders, not herbivores. The greatest impact on the monitored plants in South Korea was by the leaf beetleG. birmanicawhich defoliated most of the plants, causing the mats of plants to sink. In Europe, a similar and related insect fauna was found, but none of the insects was very damaging to the plants. One Italian weevil,Bagous rufimanus,feeds within the fruit stems and might be more damaging at higher than observed population levels. Some diseases were seen on some plants in both regions. European populations ofT. natanshave declined significantly, stimulating conservation efforts. Literature records ofTrapainsects from warm areas are presented. These could be of interest ifT. natansspreads into warmer areas of North America.     

Biological control ofSenecio jacobaea in Northern California,an enduring success

Senecio jacobaea, a poisonous weed from Eurasia, was brought under successful biological control in the Ft. Bragg, California area by 1976, through the combined action of the defoliating cinnabar moth (Tyria jacobaeae) and a root feeding flea beetle (Longitarsus jacobaeae). In 1987, 4 previously infested Ft. Bragg sites (3 sites where control had been documented and another unstudied site) were examined.Senecio jacobaea densities at these sites were 0.0, 0.0, 0.01 and 0.18 plants/m², indicating both continued and improved control of the weed. The flea beetle and the cinnabar moth both persist at the sites, despite very low numbers ofS. jacobaea plants. The control ofS. jacobaea has resulted in the return of near natural vegetation at the 2 coastal prairie sites and regained productivity at the 2 pasture sites. c/o American Embassy APO San Francisco 96301     

Host specificity ofAcanthoscelides quadridentatus andA. Puniceus [Col.: Bruchidae] for biological control ofMimosa pigra (with preliminary data on their biology)

The biology and host specificity of the seed-feeding bruchids,Acanthoscelides quadridentatus (Schaeffer) andA. puniceus Johnson, from Mexico were studied in quarantine facilities in Australia. Distinguishing characters in the genitalia of each species are illustrated. Oviposition was recorded on 16 of 73 species of plants tested but larvae died without entering pods or seeds, except onMimosa invisa K. F. P. von Martius andM. pigra L. Larvae entering seeds ofM. invisa died in the first instar. The bruchids were clearly specific toM. pigra and were subsequently released as part of a program for biological control of this weed in the Northern Territory, Australia in April 1983 and thailand in July 1984.      

Effects of stolon severing on the expansion of Alternanthera philoxeroides from terrestrial to contaminated aquatic habitats

Few studies have examined the effects of clonal integration (translocation of resources between interconnected ramets) during the expansion of amphibious clonal plants from terrestrial to aquatic habitats. We conducted a greenhouse experiment to simulate the expansion of plants from terrestrial to contaminated aquatic habitats in the amphibious stoloniferous herb Alternanthera philoxeroides (alligator weed). The proximal ramets (i.e. relatively old) of clonal fragments grown in uncontaminated soils were connected to (allowing clonal integration) or disconnected from (preventing clonal integration) distal ramets (i.e. relatively young) grown either in uncontaminated water (control, no CuSO₄) or in four copper‐contaminated water treatments containing 31.25, 62.5, 125 and 250 mg/L CuSO₄, respectively. When a stolon connection was severed, all distal ramets grown in the contaminated water died. When the stolon connection was intact, however, the survival rate of the distal ramets was 85–100% when they were grown at the three lower levels of contamination and 43.75% at the highest level. Moreover, the survival rate and growth of the distal ramets grown in the three lower levels of contamination treatments did not differ from those in the control (uncontaminated water). These results suggest that clonal integration could greatly improve the survival and growth of alligator weed subjected to moderate levels of copper stress. Although clonal integration could also increase the survival rate of the connected distal ramets subjected to the highest level of copper stress (250 mg/L CuSO₄) compared with that of disconnected distal ramets, the survival rate and growth measures were still significantly lower than those in the control. This suggests that clonal integration plays a limited role in the survival and growth of alligator weed when it is subjected to severe stress by high levels of copper contamination.     

Disturbance-mediated competition: the interacting roles of inundation regime and mechanical and herbicidal control in determining native and invasive plant abundance

Disturbance is a key component of many successful plant invasions. However, interactions among natural and anthropogenic disturbances and effects of these interacting disturbances on invasive plants and desired vegetation are rarely examined. We investigated the effect of anthropogenic disturbance (herbicidal and mechanical) along a natural inundation gradient (20–282 days) on the biomass and resource allocation of the invasive wetland plant, alligator weed (Alternanthera philoxeroides), and two co-occurring competitor plants, the introduced grass, kikuyu (Pennisetum clandestinum), and the native grass, couch (Cynodon dactylon), over a 2-year period. In the absence of additional disturbance, kikuyu biomass was negatively affected, alligator weed biomass was positively affected, and couch biomass was not affected by inundation disturbance. In addition, kikuyu was not affected by the selective removal of alligator weed, while couch increased in wetter habitats where kikuyu was absent due to inundation stress. This suggests that kikuyu is a superior competitor in drier habitats and inundation facilitates the invasion of alligator weed, while couch is an inferior competitor to both kikuyu and alligator weed and is therefore suppressed across its entire niche by these two introduced species. Mowing alone had no effect on the biomass of the species, suggesting the plants are equally tolerant of shoot removal. Selective herbicide reduced alligator weed biomass by 97.5% and the combination of selective herbicide and mowing reduced the biomass of alligator weed significantly more than herbicide alone, by 98.6% compared with un-manipulated controls. To predict community change and prevent sequential exotic plant invasions after weed removal, it is necessary to consider the interacting effects of disturbance and the niche space of invasive species in the local propagule pool.     

Clonal integration supports the expansion from terrestrial to aquatic environments of the amphibious stoloniferous herb Alternanthera philoxeroides

Effects of clonal integration on land plants have been extensively studied, but little is known about the role in amphibious plants that expand from terrestrial to aquatic conditions. We simulated expansion from terrestrial to aquatic habitats in the amphibious stoloniferous alien invasive alligator weed ( Alternanthera philoxeroides ) by growing basal ramets of clonal fragments in soils connected (allowing integration) or disconnected (preventing integration) to the apical ramets of the same fragments submerged in water to a depth of 0, 5, 10 or 15 cm. Clonal integration significantly increased growth and clonal reproduction of the apical ramets, but decreased both of these characteristics in basal ramets. Consequently, integration did not affect the performance of whole clonal fragments. We propose that alligator weed possesses a double-edged mechanism during population expansion: apical ramets in aquatic habitats can increase growth through connected basal parts in terrestrial habitats; however, once stolon connections with apical ramets are lost by external disturbance, the basal ramets in terrestrial habitats increase stolon and ramet production for rapid spreading. This may contribute greatly to the invasiveness of alligator weed and also make it very adaptable to habitats with heavy disturbance and/or highly heterogeneous resource supply.     

Introduction, Rearing, and Host Range of Aerenicopsis championi Bates (Coleoptera: Cerambycidae) for the Biological Control of Lantana camara L. in Australia

The biology and host range of the cerambycid beetle Aerenicopsis championi Bates, a potential biological control agent for the weed Lantana camara L., were studied. A. championi is a univoltine species associated with L. camara, L. urticifolia, and L. hirsuta in Mexico and Central America. In Mexico, adult emergence occurred in May and June at the start of the rainy season. Larvae fed within the stems over a 9- to 12-month period and caused damage to the plant. The insect was imported into Australia, where a procedure for rearing it in the laboratory was developed. Host-range tests indicated that adults oviposited and larvae commenced development in L. camara and L. montevidensis but not in any of 57 other species tested. A CLIMEX model indicated that most areas infested with lantana in Australia would have a favorable climate for A. championi. Permission to release this insect in Australia was obtained and three small releases were made in southern Queensland in February 1995.     

Host-specificity and biology ofMetallactus patagonicus [Col.: Chrysomelidae] introduced into Australia for the biological control ofBaccharis halimifolia [Compositae]

Biology and host-specificity of the foliage-feeding chrysomelidMetallactus patagonicus Suffrian were studied in Brazil to determine its suitability for introduction into Australia to control the shrubBaccharis halimifolia L. Multiple-choice host preference testing of plants related toBaccharis and of desirable plants from a range of plant families, showed thatM. patagonicus was restricted toBaccharis spp. It was released in Australia in 1975, but did not establish.      

Agonosoma trilineatum (Heteroptera: Scutelleridae) a biological control agent of the weed bellyache bush,Jatropha gossypiifolia (Euphorbiaceae)

Bellyache bush, Jatropha gossypiifolia L., is a serious weed of northern Australia. Agonosoma trilineatum (F.) is an insect from tropical America released in Australia in 2003 as a biological control agent against bellyache bush. It feeds on seeds and has the potential to reduce seed production, thereby potentially reducing the rate of spread and recruitment. To test the host specificity of A. trilineatum, four biological responses to host plant species were determined: development of nymphs, oviposition preferences, adult feeding and frequency of mating. Development of nymphs to adults and adult feeding only occurred on three Jatropha spp. These species also supported mating and oogenesis but only J. gossypiifolia was accepted for oviposition. Mating did not occur in the presence of other plant species. The evidence indicates that there is little risk associated with the release of this insect species in Australia and probably other countries where this weed is a problem. The probability of this insect expanding its host range is low because multiple aspects of the biology would need to change simultaneously. A. trilineatum was released in Australia between 2003 and 2007. A Climex model indicated that coastal areas of Queensland and the Northern Territory would be climatically most suitable for this insect.     

Life cycle and host specificity ofEutinobothrus sp. (Col.: curculionidae) an agent for biological control ofSida acuta (malvaceae) in the northern territory,Australia

The biology of an undescribed weevil,Eutinobothrus sp., and its specificity toSida acuta, a serious weed in northern Australia, were studied in a quarantine facility in Australia. Sixty-eight plant species were screened and onlyS. acuta andS. rhombifolia could sustain populations ofEutinobothrus sp. Adults feed externally on the stems and oviposit in feeding scars. Larvae feed internally and pupate in the stem. Prepupae and pupae may aestivate in the stem and adults emerge with the onset of the wet season when new growth is evident.Eutinobothrus sp. was released in January, 1994 in the Northern Territory and has established.     

Integration of biological control agents with other weed management technologies: Successes from the leafy spurge (Euphorbia esula) IPM program

An invasive weed can occupy a variety of environments and ecological niches and generally no single control method can be used across all areas the weed is found. Biological control agents integrated with other methods can increase and/or improve site-specific weed control, but such combinatorial approaches have not been widely utilized. The successful leafy spurge (Euphorbia esula L.) control program provides examples for future integrated weed programs that utilize biological control agents with traditional methods. Weed control methods can be used separately, such as when the leafy spurge gall midge (Spurgia esulae Gagné) reduced seed production in wooded areas while herbicides prevented further spread outside the tree line. Traditional methods also can be used directly with biological control agents. Incorporation of Aphthona spp. with herbicides has resulted in more rapid and complete leafy spurge control than either method used alone. Also, the insect population often increased rapidly following herbicide treatment, especially in areas where Aphthona spp. were established for several years but had been ineffective. Incorporation of Aphthona spp. with sheep or goat grazing has resulted in a larger decline in leafy spurge production than insects alone and in weed density than grazing alone. Controlled burns can aid establishment of biological control agents in marginally suitable environments, but timing of the fire must be coordinated to the insect’s life-cycle to ensure survival. Integration of biological control agents with revegetation programs required the agent to be the last method introduced because the cultivation and herbicide treatments necessary to establish desirable grasses and forbs were destructive to the insect. In a practical application, herbicides were combined with Aphthona spp. to help the insect establish and control leafy spurge in the habitat of the western prairie fringed orchid (Platanthera praeclara Sheviak and Bowles), an endangered species. Several experimental designs can be used to evaluate biological control agents with cultural, mechanical, and chemical control methods or with additional biological agents.     

Decomposition dynamics of invasive alligator weed compared with native sedges in a Northland lake

Invasive weeds have been shown to alter ecosystem processes such as decomposition and nutrient cycling. However, little is known about the effects of introduced biocontrol agents on these processes. This study examined the effects of alligator weed (Alternanthera philoxeroides) and its biocontrol agent, the alligator weed flea beetle (Agasicles hygrophila), on nutrient cycling in a northern New Zealand lake. Alligator weed litter decomposed significantly faster than either of two native sedge species (Schoenoplectus tabernaemontani, Isolepis prolifer) in a litterbag experiment. In addition, the presence of the alligator weed flea beetle resulted in large amounts of decaying alligator weed litter entering the lake in early summer. Both the timing and magnitude of this litter input were uncharacteristic of seasonal biomass dynamics of the native sedges. Combined with alligator weed?s rapid decomposition, this indicates altered patterns of nutrient cycling at the lake, with potential flow-on effects including facilitation of further weed invasion.