Indirect interaction between two native thistles mediated by an invasive exotic floral herbivore

Spatial and temporal variation in insect floral herbivory is common and often important. Yet, the determinants of such variation remain incompletely understood. Using 12 years of flowering data and 4 years of biweekly insect counts, we evaluated four hypotheses to explain variation in damage by the Eurasian flower head weevil, Rhinocyllus conicus, to the native North American wavyleaf thistle, Cirsium undulatum. The four factors hypothesized to influence weevil impact were variations in climate, weevil abundance, phenological synchrony, and number of flower heads available, either on wavyleaf thistle or on the other cooccurring, acquired native host plant (Platte thistle, Cirsium canescens), or on both. Climate did not contribute significantly to an explanation of variation in R. conicus damage to wavyleaf thistle. However, climate did influence weevil synchrony with wavyleaf flower head initiation, and phenological synchrony was important in determining R. conicus oviposition levels on wavyleaf thistle. The earlier R. conicus was active, the less it oviposited on wavyleaf thistle, even when weevils were abundant. Neither weevil abundance nor availability of wavyleaf flower heads predicted R. conicus egg load. Instead, the strongest predictor of R. conicus egg load on wavyleaf thistle was the availability of flower heads on Platte thistle, the more common, earlier flowering native thistle in the sand prairie. Egg load on wavyleaf thistle decreased as the number of Platte thistle flower heads at a site increased. Thus, wavyleaf thistle experienced associational defense in the presence of flowering by its now declining native congener, Platte thistle. These results demonstrate that prediction of damage to a native plant by an exotic insect may require knowledge of both likely phenological synchrony and total resource availability to the herbivore, including resources provided by other nontarget native species.     

Phenological synchrony affects interaction strength of an exotic weevil with Platte thistle,a native host plant

Phenological synchrony of a consumer population with its resource populations is expected to affect interaction intensity. We quantified phenological variation and synchrony of populations of an invasive Eurasian flower head weevil, Rhinocyllus conicus, that consumes florets, ovules, and seeds of developing flower heads of a native North American thistle, Cirsium canescens, in Sand Hills prairie in Nebraska, USA. Variation in timing of adult activity among weevil populations was larger than variation in timing of flower head development among C. canescens populations, and it drove the observed variation in the phenological synchrony between weevil and host plant populations. Furthermore, the degree of phenological synchrony between populations was significant in explaining variation in weevil egg load on the newly acquired host plant. Because population growth of C. canescens is limited by predispersal seed losses to floral herbivores, variation in the synchrony of herbivore and plant flowering will affect the density of the plant population. These results provide strong quantitative support for the hypothesis that the synchrony of insect activity with plant resources can determine the magnitude of impact of floral herbivores on their host plant populations.     

Exotic weevil invasion increases floral herbivore community density,function, and impact on a native plant

Consumer communities are being re‐arranged through unprecedented rates of human‐mediated invasions and extinctions. Such changes in consumer diversity potentially alter community function and impact on resource populations. Although insect herbivore invasions are increasingly common, the influence of such species additions on native resident herbivore guilds, along with their individual and combined effects on native plant resources, are rarely investigated. Here, we used site‐to‐site and plant‐to‐plant variation in herbivore composition to examine how the addition of an invasive exotic weevil, Rhinocyllus conicus, combines with a guild of native floral herbivores (tephritid flies, pyralid moths) to influence two key components of herbivore community function – aggregate herbivore densities and cumulative levels of seed destruction – on a native thistle, Cirsium canescens. Invasion of a site by R. conicus more than doubled aggregate herbivore density, resulting in increased levels of seed destruction and a halving of seed production by the native thistle. Further, herbivore function was significantly higher on individual plants attacked by R. conicus, compared to plants attacked only by native herbivores. Insect densities and levels of seed destruction on plants attacked by multiple herbivore taxa never exceeded those observed for plants attacked by R. conicus alone, suggesting that increases in herbivore community function with invasion resulted from the inclusion of a functionally dominant insect rather than any complementarity effects. Some evidence for interference between insects emerged, with a trend towards reduced moth and weevil densities in two and three taxon mixtures compared with plants attacked by each taxon alone. However, density compensation was limited so that, overall, the addition of a novel herbivore to the floral guild was associated with a significant increase in herbivore community function and impact on seed production. The results suggest that invasion of a functionally dominant herbivore into an unsaturated recipient community can augment function within a resource guild.     

The invasive <Emphasis Type="Italic">Lespedeza cuneata</Emphasis> attracts more insect pollinators than native congeners in tallgrass prairie with variable impacts

Invasive plant species can potentially exert competitive or facilitative effects on insect pollination services of native species. Factors that influence these effects include the degree of shared pollinator species, synchronous flowering phenology, similar flower morphology and color, relatedness of invasive and natives, and showiness and densities of flowers. We investigated such plant-pollinator dynamics by comparing the invasive Lespedeza cuneata and three native congeners, all sympatric with synchronous flowering, using in situ populations over 2 years during peak floral displays. Insect visitation rates of the invasive were significantly higher per plant in both years than on the native species. The invasive exerted a competitive effect on visitation of the two native species with fewer shared pollinators, and a facilitative effect on visitation of the native species with the highest degree of shared insect visitors. Positive correlations were found between floral density and visitation rate per plant in all the native species. Although no such correlation was found for the invasive, floral density in L. cuneata was at least twenty times higher than in the native species and likely saturated the response of the pollinator community. Analyses of insect visitor taxonomic data indicated the insect communities visiting each of the Lespedeza species were generally similar though with species-specific differences. The main exception was that the common honeybee, Apis mellifera, was a primary visitor to the invasive plant species, yet was never observed on the native Lespedeza species.     

Individually mark–mass release–resight study elucidates effects of patch characteristics and distance on host patch location by an insect herbivore

1. How organisms locate their hosts is of fundamental importance in a variety of basic and applied ecological fields, including population dynamics, invasive species management and biological control. However, tracking movement of small organisms, such as insects, poses significant logistical challenges. 2. Mass‐release and individual–mark–recapture techniques were combined in an individually mark–mass release–resight (IMMRR) approach to track the movement of over 2000 adult insects in an economically important plant–herbivore system. Despite its widespread use for the biological control of the invasive thistle Carduus nutans, the host‐finding behaviour of the thistle head weevil Rhinocyllus conicus has not previously been studied. Insects were released at different distances from a mosaic of artificially created host patches with different areas and number of plants to assess the ecological determinants of patch finding. 3. The study was able to characterize the within‐season dispersal abilities and between‐patch movement patterns of R. conicus. Weevils found host plant patches over 900 m away. Large patches, with tall plants, situated close to the nearest release point had the highest first R. conicus resights. Patch area and plant density had no effect on the number of weevils resighted per plant; however, R. conicus individuals were more likely to disperse out of small patches and into large patches. 4. By understanding how R. conicus locates host patches of C. nutans, management activities for the control of this invasive thistle can be better informed. A deeper mechanistic understanding of host location will also improve prediction of coupled plant–herbivore spatial dynamics in general.     

Host recognition by Rhinocyllus conicus of floral scents from invasive and threatened thistles

One of the main obstacles of classical biological control is that biological control organisms cannot be recalled once they are released in nature. It is particularly true for the flowerhead weevil, Rhinocyllus conicus Frölich, which was released as a biological control organism for the invasive musk thistle, Carduus nutans L. (MT). While weevils successfully suppressed introduced populations of musk thistles and other invasive thistle species, non-target attacks have been reported on multiple native thistles including federally listed threatened and endangered (T&E) thistle species. To investigate the foraging behavior of female weevils on invasive and native thistles, we examined volatile organic compounds (VOCs) emitted from MT and a T&E plant species, Sacramento Mountains thistle, Cirsium vinaceum Wooton & Standley (SMT) in the Lincoln National Forest, New Mexico. We used a dynamic headspace volatile collection system and gas chromatography-mass spectrometry to compare volatile profiles between MT and SMT. Female weevils reacted to 7 electrophysiologically active chemical compounds in the blends based on gas chromatography-electroantennography. The behavioral response of female weevils was indifferent when VOCs from both thistles were offered in y-tube olfactometry experiments. Yet, they preferred VOCs collected from MT to purified air. The searching time of female weevils was longer to VOCs collected from SMT over controls. Investigating signals during the initial host recognition of released biological control organisms may open new opportunities to reduce non-target attacks on T&E plant species.

     

Interference and exploitation competition of three nectar-thieving invasive ant species

Summary. Plant and insect exudates are known to play a key role in structuring tropical ant communities, but less is known about the utilization of these resources in communities dominated by invasive ants. Invasive ants are thought to require large amounts of carbohydrates such as honeydew or nectar to maintain their high abundances. Invasive ants that consume floral nectar may compete with legitimate floral visitors through interference or exploitation competition. I compared the nectar-thieving behavior of three widespread invasive ant species: long-legged ants (Anoplolepis gracilipes), Argentine ants (Linepithema humile), and big-headed ants (Pheidole megacephala) in inflorescences of the native Hawaiian ‘ōhi’a tree, an important food source for native fauna. A. gracilipes was least likely to leave inflorescences unvisited and visited inflorescences in higher numbers than both L. humile and P. megacephala. A. gracilipes and L. humile visited more flowers in an inflorescence and were less likely to retreat from a flower with a competitor than P. megacephala. A. gracilipes was able to take 5.5 and 11.3 times the amount of nectar than L. humile and P. megacephala, respectively. Thus, A. gracilipes may be effective at both interference and exploitation competition against other nectarivores, L. humile may be effective at interference competition, and P. megacephala may be relatively weak at both types of competition against other nectarivores. Ascertaining the competitive abilities of invasive ants against legitimate floral visitors will be especially important in agricultural and other systems that are nectar or pollinator limited.Received 6 December 2004; revised 13 January 2005; accepted 14 January 2005.     

Effects of Rhinoncomimus latipes on the growth and reproduction of Persicaria perfoliata,an invasive plant in North America

Native to Asia, mile-a-minute Persicaria perfoliata, is an invasive weed in North America, and the weevil Rhinoncomimus latipes is a host-specific insect agent which occurs widely in China. We conducted a common garden experiment to compare P. perfoliata plant responses of native and invasive populations to herbivory by the weevils from different origins. We found weevils from Hunan, Hubei and Heilongjiang Provinces had strong, moderate and weak ability to suppress host plant, respectively. Weevils from Hunan and Hubei Provinces had stronger impact on the growth of both native and invasive plant populations than the weevil from Heilongjiang Province. The losses in seed output of invasive plants were also significantly greater than natives in the weevil treatments. Our results suggested that the weevil population from Hunan Province may be the most suitable for the control of mile-a-minute, while the population from Heilongjiang Province may be the least suitable due to climate matching.     

Non-target host risk assessment of the idiobiont parasitoid Bracon celer (Hymenoptera: Braconidae) for biological control of olive fruit fly in California

Non-target risk posed by an African parasitoid, Bracon celer Szépligeti (Hymenoptera: Braconidae), was assessed for a classical biological control program against olive fruit fly, Bactrocera oleae (Rossi) (Diptera: Tephritidae: Dacinae), in California, USA. Behavioral and reproductive responses to non-target tephritid species were tested with beneficial (Chaetorellia succinea [Costa] and Parafreutreta regalis Munro) (Tephritidae: Tephritinae) and native (Rhagoletis fausta [Osten Sacken]) (Tephritidae: Trypetinae) fruit fly species in successive no-choice and choice experiments under close confinement in quarantine. Non-target host-plant substrates exposed to B. celer were yellow-starthistle flower heads containing C. succinea, Cape ivy stem galls containing P. regalis, and bitter-cherry fruit containing R. fausta. The parasitoid probed all three infested non-target plant substrates, but significantly less than olives infested with B. oleae. It produced offspring from P. regalis in Cape ivy stem galls, but appeared unable to penetrate yellow-starthistle flower heads with its ovipositor. Bracon celer killed some B. oleae and R. fausta larvae without parasitism. Reproduction on P. regalis indicates that B. celer has a broad physiological host range, which, combined with the parasitoid's acceptance of all three host-plant substrates, indicates a strong potential to negatively impact non-target species. Although physical and temporal barriers to host attack may reduce risk to most non-target tephritids by B. celer in California, the parasitoid should not be released due to its risk of harming the beneficial P. regalis. Release of P. regalis is still under consideration, however, and final risk assessment should depend on whether the fly proves useful for weed control.     

Consequences of variation in flowering phenology for seed head herbivory and reproductive success in Erigeron glaucus (Compositae)

Summary We examined the relationship between flowering phenology, reproductive success (seed production only), and seed head herbivory for 20 similarly sized clones of Erigeron glaucus growing at Bodega Bay Reserve, northern California, USA. Although clones tended to reach peak flowering on the same date, they differed in the proportion of their total flowers produced around that date (flowering synchrony). Clones also differed in the number and density of flower heads presented at any one time to pollinators and herbivores (floral display). Both of these characteristics had consequences for herbivory and plant reproductive success. The proportion of flower heads damaged by insect herbivores was greater for clones that concentrated flowering activity during the main flowering period for the population as a whole (high synchrony) compared to clones that spread flowering out temporally. The primary reason for this result was that clones with low flowering synchrony produced a significant proportion of their flower heads during the fall and therefore, escaped attack by the tephritid fly, Tephritis ovatipennis. Clones with intermediate synchrony had lower seed success (total number of viable seeds produced over the year) than clones with either low or high synchrony. The proportion of flower heads damaged by insect herbivores and number of tephritid flies reared from flower heads were both negatively correlated to floral display while seed head mass and germination rates were positively related to display. Thus, clones which produced dense floral displays were favored both in terms of reduced herbivory and increased successful seed production.     

Inter-annual variation in above- and belowground herbivory on a native,annual legume

The relative importance of subterranean versus aboveground insect damage to plants is not well understood. In particular, the simultaneous effects of above- and belowground herbivory, and the importance of highly variable abiotic factors such as rainfall, have received little attention in diverse natural ecosystems. We investigated the influence of both above- and belowground herbivory on Lupinus nanus (Fabaceae), an annual plant native to coastal California. A number of insect species damage L. nanus aboveground, and a weevil larva consumes nodules belowground. To manipulate herbivory in the field, we employed a combination of insecticides and simulated herbivory during two different years. In 1997, simulated belowground damage reduced L. nanus survival, and insecticide application to roots increased seed production and seed mass. By contrast, in 1998, only aboveground folivory significantly reduced L. nanus reproduction, and, in combination, above- and belowground insecticides did not affect flower or seed number relative to controls. A growth chamber experiment conducted in the absence of herbivory revealed that the aboveground insecticide marginally reduced flower production and the belowground insecticide marginally increased flower production compared to controls; these non-target effects made our field experiments for aboveground herbivory conservative. Finally, ambient levels of herbivory differed among years (1997, 1998, and 2000), which varied greatly in rainfall due to the effects of El Nino. The results suggest that the impacts of herbivores are temporally variable and that abiotic factors, particularly those related to large-scale changes in weather patterns, may be more important than insect herbivory to L. nanus in some years. This revised version was published online in August 2006 with corrections to the Cover Date.     

Should the flower bud weevil Anthonomus santacruzi (Coleoptera: Curculionidae) be considered for release against the invasive tree Solanum mauritianum (Solanaceae) in New Zealand?

The invasive tree Solanum mauritianum Scop. has been targeted for biological control in South Africa and New Zealand, by deploying insect agents that could constrain its excessive reproductive output. The flower-feeding weevil Anthonomus santacruzi (Curculionidae) was approved for release in South Africa in 2007 but following the loss of the original culture in quarantine, new stocks were introduced from Argentina in 2008–2009. This study was initiated to confirm that the host range of the new culture was the same as that of the previous one, but also to assess the risks associated with the weevil's release in New Zealand. Different testing procedures, including no-choice tests and multi-choice tests in different arenas, produced inconsistent and ambiguous results. During no-choice tests, oviposition and larval development to adulthood occurred on three non-target species including two native South African and one native New Zealand Solanum species. However, subsequent multi-choice tests and a risk assessment suggested that the risks of anything more than collateral damage to non-target Solanum species are low. Overall, these data do not deviate substantially from the results of the original quarantine tests which facilitated the release of A. santacruzi in South Africa in 2009. Although we argue that none of the New Zealand native and cultivated species are at risk, stronger evidence from open-field trials and chemical ecology studies may be required to convince the regulatory authorities that A. santacruzi is suitable for release in New Zealand.     

Biotic resistance and invasional meltdown: consequences of acquired interspecific interactions for an invasive orchid,Spathoglottis plicata in Puerto Rico

Invasiveness of non-native species often depends on acquired interactions with either native or naturalized species. A natural colonizer, the autogamous, invasive orchid Spathoglottis plicata has acquired at least three interspecific interactions in Puerto Rico: a mycorrhizal fungus essential for seed germination and early development; a native, orchid-specialist weevil, Stethobaris polita, which eats perianth parts and oviposits in developing fruits; and ants, primarily invasive Solenopsis invicta, that forage at extrafloral nectaries. We tested in field experiments and from observational data whether weevils affect reproductive success in the orchid; and whether this interaction is density-dependent. We also examined the effectiveness of extrafloral nectaries in attracting ants that ward off weevils. Only at small spatial scales were weevil abundance and flower damage correlated with flower densities. Plants protected from weevils had less floral damage and higher fruit set than those accessible to weevils. The more abundant ants were on inflorescences, the less accessible fruits were to weevils, resulting in reduced fruit loss from larval infections. Ants did not exclude weevils, but they affected weevil activity. Native herbivores generally provide some biotic resistance to plant invasions yet Spathoglottis plicata remains an aggressive colonizer despite the acquisition of a herbivore/seed predator partly because invasive ants attracted to extrafloral nectaries inhibited weevil behavior. Thus, the invasion of one species facilitates the success of another as in invasional meltdowns. For invasive plant species of disturbed habitats, having ant-tended extrafloral nectaries and producing copious quantities of seed, biotic resistance to plant invasions can be minimal.     

Physiological host range of two highly specialised mutualistic symbiotes: the fly Fergusonina turneri and its obligate nematode Fergusobia quinquenerviae,potential biocontrol agents of Melaleuca quinquenervia

In Australia, galls develop on Melaleuca quinquenervia (Cav.) S.T. Blake (Myrtaceae) as a result of the mutualistic association between the fly Fergusonina turneri Taylor (Diptera: Fergusoninidae) and its obligate nematode Fergusobia quinquenerviae Davies & Giblin-Davis (Tylenchida: Sphaerulariidae). The nematode induces gall formation, whereas the fly promotes gall maturation. Together they exploit M. quinquenervia buds and may inhibit stem elongation and flower formation. We delimited the physiological host range of this pair to determine their suitability as biological control agents of invasive M. quinquenervia populations in Florida, USA. Host use was assessed for eight species of Myrtaceae native to Florida, eight phylogenetically related ornamental species and oviposition alone on five non-myrtaceous species. Although oviposition was less specific, galls developed and matured only on M. quinquenervia. After establishment, galls are predicted to prevent flower and seed production, thereby reducing the regenerative potential of M. quinquenervia. This is the first example of an insect/nematode mutualism released as biological control agents of an invasive plant.     

Insect-Flower Interaction Network Structure Is Resilient to a Temporary Pulse of Floral Resources from Invasive Rhododendron ponticum

Invasive alien plants can compete with native plants for resources, and may ultimately decrease native plant diversity and/or abundance in invaded sites. This could have consequences for native mutualistic interactions, such as pollination. Although invasive plants often become highly connected in plant-pollinator interaction networks, in temperate climates they usually only flower for part of the season. Unless sufficient alternative plants flower outside this period, whole-season floral resources may be reduced by invasion. We hypothesized that the cessation of flowering of a dominant invasive plant would lead to dramatic, seasonal compositional changes in plant-pollinator communities, and subsequent changes in network structure. We investigated variation in floral resources, flower-visiting insect communities, and interaction networks during and after the flowering of invasive Rhododendron ponticum in four invaded Irish woodland sites. Floral resources decreased significantly after R. ponticum flowering, but the magnitude of the decrease varied among sites. Neither insect abundance nor richness varied between the two periods (during and after R. ponticum flowering), yet insect community composition was distinct, mostly due to a significant reduction in Bombus abundance after flowering. During flowering R. ponticum was frequently visited by Bombus; after flowering, these highly mobile pollinators presumably left to find alternative floral resources. Despite compositional changes, however, network structural properties remained stable after R. ponticum flowering ceased: generality increased, but quantitative connectance, interaction evenness, vulnerability, H’₂ and network size did not change. This is likely because after R. ponticum flowering, two to three alternative plant species became prominent in networks and insects increased their diet breadth, as indicated by the increase in network-level generality. We conclude that network structure is robust to seasonal changes in floral abundance at sites invaded by alien, mass-flowering plant species, as long as alternative floral resources remain throughout the season to support the flower-visiting community.     

Sex‐biased seed predation in gynodioecious Dianthus superbus var. longicalycinus (Capryophyllaceae) and differential influence of two seed predator species on the floral traits

Gynodioecy, the co‐occurrence of hermaphrodite and female individuals within a species, is maintained by differential reproductive success between sexes. Recently, researchers recognized that not only pollinators but also herbivores are important agents in the evolution and maintenance of gynodioecy, when herbivory is hermaphrodite biased. In this study, we investigated whether there is hermaphrodite‐biased herbivory in a gynodioecious plant, Dianthus superbus var. longicalycinus, and if so, what floral traits influenced hermaphrodite‐biased herbivory. We measured flower morphology (flower diameter, calyx tube length, corolla height and petal width) and phenology of flowers of female individuals, hermaphrodites and gynomonoecious individuals in a natural population. We also investigated seed predation and predator species. At the study site, Sibinia weevils (Curculionidae; Coleoptera) and Coleophora moths (Coleophoridae; Lepidoptera) were common pre‐dispersal seed predators in this species. The weevil appeared early in the flowering season, and weevil predation correlated with flower phenology. Because female individuals did not flower early in the season, weevil predation was less frequent in female individuals. Moth predation correlated with calyx length. The calyx length of flowers of female individuals was smaller than those of hermaphrodites, but a direct comparison of moth predation rates failed to find a significant difference among sex morphs. We found that the two seed predators had different effects on floral traits in D. superbus var. longicalycinus. We suggest that weevil predation contributes to the maintenance of gynodioecy because female individuals successfully escaped weevil predation by flowering late. It remains unclear why flower phenology is different among sex morphs.     

Timing of reproduction in a prairie legume: seasonal impacts of insects consuming flowers and seeds

Summary Seasonal patterns of insect damage to reproductive tissue of the legume Baptisia australis were studied for three years in native tallgrass priairie. Contrasting seasonal patterns of damage were associated with the major species of insect consumers. The moth Grapholitha tristegana (Olethreutidae) and the weevil Tychius sordidus (Curculionidae), which together infested 80–100% of developing fruits (pods), consistently damaged more seeds on average in early than in late maturing pods. But while late opening flowers were less subject to attack from moths and weevils, they were more subject to attack from chewing insects, particularly blister beetles (Epicauta fabricii, Meloidae), which destroyed >80% of all flowers and developing young pods (including moth and weevil larval inhabitants). The blister beetles arrived late in the flowering season and fed particularly on young reproductive tissue, allowing larger, older pods that had developed from early opening flowers to escape destruction. The relative abundances and impacts of blister beetles, moths, and weevils varied from year to year. Adding to the uncertainty of reproductive success of the host plant were the large and variable amounts of damage to immature buds inflicted by insects (including the blister beetles and weevil adults) and late killing frosts. Thus, timing of flowering is critical to success in seed production for B. australis. The heavy impacts of insects and weather can result in a very narrow window in time (which shifts from year to year) during which B. australis can flower with any success. The opposing pressures exerted by insects and weather on floral reproductive success may act in concert with other features of the plant's biology to foster the maintenance of considerable diversity in flowering times among individuals in local populations of B. australis.     

Comparison of nectar use and preference in the parasitoid Trybliographa rapae (Hymenoptera: Figitidae) and its host,the cabbage root fly,Delia radicum (Diptera: Anthomyiidae)

This study investigated differences in flower preferences between the parasitoid Trybliographa rapae Westwood (Hymenoptera: Figitidae) and its host, the economically important pest of cruciferous crops, the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The data obtained were used to suggest selective food plants in conservation biological control programmes for control of D. radicum. The attraction of both insect species to floral odours emitted from nine different plant species, their ability to access nectar from four of these species and the effect of the most promising plant species on insect longevity were determined. Naive T. rapae females were significantly attracted to flower odours from Fagopyrum esculentum Moench. (Polygonaceae) and repelled by Coriandrum sativum L. (Apiaceae) and Borago officinalis L. (Boraginaceae). In addition, T. rapae gained weight when exposed to F. esculentum, Anethum graveolens L. (Apiaceae) and Lobularia maritima Desv. (Brassicaceae). In contrast, naive D. radicum females showed attraction to most of the flowers. The longevity of both T. rapae and D. radicum increased significantly when they were provided with flowering A. graveolens and F. esculentum. In addition to the laboratory studies, a semi-field experiment was made to study the impact of flowering F. esculentum on the ability of T. rapae to parasitise D. radicum larvae. Significantly more larvae were parasitised in cages where a floral resource was present. The findings are discussed in the context of a Brassica agroecosystem.     

Simulated herbivory counteracts the effects of grass competition on the invasive fireweed (Senecio madagascariensis)

Competition, herbivory and their interaction play a significant role in determining the competitive ability and survival of individual plant species. Understanding these processes and interactions can improve the efficacy of biocontrol programs against invasive weeds. Senecio madagascariensis (fireweed) is an invasive weed of South African origin that reduces pastoral productivity and poisons livestock in several countries, notably Australia. Although competitive pastures can suppress the weed’s growth in Australia, its competitive nature is poorly understood in relation to its invasion success. This greenhouse study assessed the growth and reproductive yield of fireweed growing in competition with six native and introduced grasses present in both South Africa and Australia. Since fireweed is a target for biocontrol in Australia, we examined whether its response to grass competition changed with herbivory (simulated by 40% leaf removal). The effect of grass competition and herbivory on the weed’s biomass and floral productivity was examined during a 12‐week pot trial in South Africa. Floral numbers were unaffected by both grass competition and herbivory. Biomass was used to calculate Relative Interaction Indices (RII) to quantify the weed’s competitive or facilitative response. This index compares a specific measurable trait, such as biomass, of fireweed growing alone, to fireweed growing with grass to determine the level of competitive suppression or facilitation resulting from the interaction. Despite the lack of species‐specific effects of grass competition, the presence of grass suppressed fireweed’s foliar, root and whole plant biomass the most when herbivory was absent. With herbivory, fireweed did not suffer from any measurable competitive suppression. This lack of competitive suppression may be due to an induced allelopathic response, given the levels of pyrrolizidine alkaloids common in many Senecio species. Since this result may weaken the case for biocontrol, the weed’s competitive responses should be verified in relation to actual insect herbivory.     

Cover Caption

Persicaria perfoliata is an invasive prickly vine in North America. The differences in enemy composition between invasive and native range may lead to new plant–insect interaction. The results of common garden experiment indicated invasive plants possess high reproduction and physical defense, and low level of defense against the specialists, especially Rhinoncomimus latipes (the weevil in the image). The high reproductive capacity and lack of effective specialist herbivores in the invasive range are the potential reasons of its invasiveness. See pages 521–532. Photo taken by Xiao‐Qiao Li.