Effect of Leafy Spurge (Euphorbia esula) Genotype on Feeding Damage and Reproduction of Aphthona spp.: Implications for Biological Weed Control

North American leafy spurge (Euphorbia esula L.) is genetically diverse and composed of multiple genotypes introduced from several areas of Europe and Asia. Five species of leafy spurge flea beetle (Aphthona spp.) have been introduced as biological control agents for leafy spurge, but were collected in a relatively small region of Europe. Greenhouse and field experiments were conducted to determine if observed variation in feeding preference and reproduction of Aphthona spp. on North American leafy spurge may be due in part to leafy spurge genotype. Leafy spurge genotypes were collected from Austria; Manitoba in Canada; and the states of Montana, Nebraska, North Dakota, South Dakota, and Wyoming in the United States. Leafy spurge genotype affected feeding but not egg laying by Aphthona spp. adults. Aphthona czwalinae/lacertosa fed slightly less in a free-choice test on a genotype from Manitoba (7%) compared to genotypes from Nebraska and North Dakota (14%). Aphthona flava tended to feed less on the Nebraska genotype than any other genotype evaluated. Reproduction of Aphthona spp. was greatly affected by leafy spurge genotype. For instance, A. czwalinae/lacertosa produced 72 adults per plant from a Nebraska genotype compared to 11 to 32 adults per plant from all other genotypes evaluated. Thus, some observed variation in establishment and reproduction of Aphthona spp. can be attributed to leafy spurge genotype.     

Temporal dynamics of leafy spurge (Euphorbia esula) and two species of flea beetles (Aphthona spp.) used as biological control agents

The goal of this study was to evaluate the biological control program of leafy spurge (Euphorbia esula) in a large natural area, Theodore Roosevelt National Park, western North Dakota, USA. Aphthona lacertosa and Aphthona nigriscutis have been released at more than 1800 points in the 18,600-ha South Unit of the park beginning in 1989; most releases have occurred since 1994. We established permanent vegetation plots throughout the infested area of the park and determined stem counts and biomass of leafy spurge and abundance of the two flea beetle species at these plots each year from 1999 to 2001. Both biomass and stem counts declined over the 3 years of the study. Both species of flea beetle are well established within the park and have expanded into areas where they were not released. A. nigriscutis was more abundant than A. lacertosa in the grassland areas we surveyed, but in all other habitats abundances were similar. Using structural equation models, only A. lacertosa could be shown to have a significant effect on counts of mature stems of leafy spurge. A. nigriscutis numbers were positively correlated with stem counts of mature stems. Previous year’s stem counts had the greatest influence on change in stem counts over each 2-year time step examined with structural equation models.     

Evidence for the influence of conspecific chemical cues on <Emphasis Type="Italic">Aphthona nigriscutis</Emphasis> (Coleoptera: Chrysomelidae) behaviour and distribution

Although the distribution of biological control agents may have a significant effect upon their impacts, the mechanisms regulating these distributions are often unknown. Such is the case with Aphthona nigriscutis, a classical biological control agent of leafy spurge in North America. These beetles assume aggregated distributions at some sites but disperse rapidly at others. The potential influence of plant and insect-factors upon aggregation and dispersal was investigated to try to explain these observations. Male beetles produce a putative aggregation pheromone. Responses of conspecifics to male-associated cues are greater when beetles are feeding on host plants. Densities of beetle groups greatly impact their attractiveness. Males are more sensitive to dispersal cues and females are more sensitive to congregation cues.     

Post hoc assessment of an operational biocontrol program: efficacy of the flea beetle Aphthona lacertosa Rosenhauer (Chrysomelidae: Coleoptera), an introduced biocontrol agent for leafy spurge

Mixed populations of Aphthona lacertosa and Aphthona czwalinae were released at more than 50 locations in Alberta in 1997. Two and 3 years post-release, beetle populations were primarily A. lacertosa, with A. czwalinae forming less than 0.5% of the sampled populations. Beetle densities were moderate (10–70 beetles per m²) or high (>70 beetles per m²) at 14% and more than 60% of the sampled sites in 1999 and 2000, respectively. Larger beetles had greater instantaneous egg loads (r²=0.424,P=0.003). In 2000, the largest beetles were found at moderate density sites and there was a significant negative relationship between beetle size and the time taken to accumulate a degree day threshold of 1230 (for females: r²=0.678,P=0.001). Sites with the most rapid accumulation of degree days have the greatest potential for beetle population growth based on potential fecundity. Changes in leafy spurge percent cover, stem density, and canopy height from 1997 to 2000 were assessed across sites with low (<10 beetles per m²), moderate, and high beetle densities in 2000. Sites with high beetle densities had significantly greater reductions of leafy spurge within 5 m of the release point than sites with low beetle densities (P<0.017). Damage caused by the beetles at high-density sites was often visible as a halo-shaped patch of dead leafy spurge stems. The significant overall reduction of leafy spurge within release patches makes A. lacertosa a promising biocontrol agent for leafy spurge in Alberta.     

Phytotoxic Allelochemicals From Roots and Root Exudates of Leafy Spurge (Euphorbia esula L.)

Invasive plants are a widespread problem but the mechanisms used by these plants to become invasive are often unknown. The production of phytotoxic natural products by invasive weeds is one mechanism by which these species may become successful competitors. Here we conducted a bioactivity-driven fractionation of root extracts and exudates from the invasive plant leafy spurge (Euphorbia esula L.), and structurally characterized jatrophane diterpenes and ellagic acid derivatives. Ellagic acid derivatives and one of the jatrophane diterpenes, esulone A, have been previously reported from leafy spurge, but another of the jatrophane diterpenes, kasuinine B, has not. We show that these compounds are phytotoxic but affect plants in different ways, either inducing overall plant necrosis or reducing root branching and elongation.Key Words: phytotoxicity, allelochemicals, roots, root exudates, jatrophane diterpenes, kansuinine B, ellagic acid derivatives, leafy spurge, Euphorbia esula, Arabidopsis thaliana     

Biology and host specificity ofChamaesphecia hungarica andCh. Astatiformis (Lep.: Sesiidae) two candidates for the biological control of leafy spurge,Euphorbia esula (Euphorbiaceae) in North America

Leafy spurge (Euphorbia esula (s.1.)) is an herbaceous perennial and serious weed of Eurasian origin that has been accidentally introduced into North America. The two European root-boring mothsChamaesphecia hungarica andCh. astatiformis are univoltine and overwinter as mature larvae. Both species have a lower survival rate on leafy spurge than on their field hosts, and thus are not optimal candidates for the biological control of leafy spurge. However, the rate of larval development and larval growth on the target weed and on the two field hosts is nearly the same. The experimental host range of both species is restricted to a few species in the subgenusEsula within the genusEuphorbia. The two species occupy different habitats in the steppe biome and are targeted for similar leafy spurge habitats in North America.     

Energy potential of leafy spurge (Euphorbia esula)

Leafy spurge (Euphorbia esula) is a noxious, perennial weed that infests pastures, rangeland and waste areas in the northern Great Plains. The objective of this study was to determine the productive potential of this species when grown under optimum agronomic conditions. Plants were fertilized and irrigated. Oil, hydrocarbon, total protein, and dry-weight production were measured on 3 harvest dates. Calorimetric analyses were performed to determine the potential of leafy spurge as a fuel crop. The hydrocarbon content of 12 strains of leafy spurge was determined to measure genetic variability for this trait. The addition of fertilizer doubled dryweight production but did not affect percent oil or hydrocarbon content. Oil and hydrocarbon production averaged 6.8 and 0.6% on a plant dry-weight basis. Maximum production of plant biomass, protein, and hydrocarbon was obtained from a mid-July harvest. Oil content increased later in the growing season. The total protein content of leafy spurge averaged 12%. Whole-plant biomass had a caloric value of 4,407 cal/g while the oils contained 10,019 cal/g. Leafy spurge hay can produce 4 times more energy per year than wheat straw; therefore, the immediate potential of leafy spurge whole-plant biomass as a locally grown fuel crop for home-heating purposes is suggested.     

Field and laboratory biology of the stem-feeding beetle Thamnurgus euphorbiae (Küster) (Col., Scolytidae) in Italy, a potential biological control candidate of leafy spurge in the USA and Canada

Thamnurgus euphorbiae overwinters as adults in dry stems of Euphorbia characias until the end of March before exiting through circular holes cut with their mandibles. Females and males appeared in the field at the same time, and the first pairs were observed on 28 March on new stems of E. characias having the open‐flower at the beginning of its development. Throughout the rest of April it was possible to find and collect the fairly abundant adults. Mating occurs on new stems of the host plant and females walk up towards the top of the stem and start to mine the centre of it in order to penetrate. Oviposition occurs in new stems and eggs are laid singly along the interior of the stem; females laid 35–85 eggs. Upon hatching, larvae started feeding on vascular bundles and the inner cortex. There are three larval instars, and pupation occurred in the stem of the host plant; this insect is univoltine. Thamnurgus euphorbiae has been accepted by the Technical Advisory Group for Biological Control Agents of Weeds for release as a biological control agent of leafy spurge in the US. Prior to this study the biology of T. euphorbiae was unknown.     

Nature,nurture, and vegetation management: Studies with sheep and goats

Diet selection and preference by grazing animals are determined by genetic and environmental factors (i.e., nature and nurture) that interact and affect their efficacy for managing vegetation as targeted grazers. The effect of rearing environment on the consumption of leafy spurge by sheep and goats was investigated. We hypothesized that although rearing environment will affect the preference for chemically defended plants ultimately, the inherent ability to detoxify or eliminate phytotoxins will limit an animal’s preference for them. The objective of this study was to determine if sheep would consume more of the invasive weed leafy spurge (Euphorbia esula) if they were raised by goat compared to sheep raised by sheep and goat raised by goat. Sheep were raised on leafy spurge-infested pastures by either their ewe (S) or a goat doe (FS) on which they were fostered within 24 hours of birth and parturition of lamb and doe, respectively. Does that fostered lambs also raised their own goat offspring (G) such that the same doe raised the FS and G animals. The rearing environment’s effect on leafy spurge consumption was tested the following growing season by simultaneously grazing all animals on the same leafy spurge-infested rangeland and estimating percentage leafy spurge in their diet with either fecal near-infrared spectroscopy (f.NIR) or bite count. Goats consumed more leafy spurge as determined by either f.NIR (62.8%, P < 0.06) or bite count (71.9%, P < 0.01) than FS (35.2 % f.NIR, 39.3% bite count) or S (10.1 % f.NIR, 18.2% bite count). The FS consumed over twice as much leafy spurge as S and were numerically intermediate to G and S for leafy spurge consumption but not significantly different from the S sheep, most likely because one FS sheep did not eat leafy spurge during the evaluation period. Because leafy spurge is aversive to sheep but not goats, higher leafy spurge consumption by FS sheep is hypothesized to result from inoculation of their rumen microbes with microbes from the does capable of denaturing aversive phytotoxins in leafy spurge. The higher consumption of leafy spurge by G compared to FS shows that genetically determined physiological differences influence an animal’s ability to ameliorate phytotoxins and determine the upper limit of an animal’s preference for a chemically defended plant. It also indicated that in addition to the animal’s genome, the genome of an animal’s microbiome, which the mother may influence, can play an important role in diet selection.     

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

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

Biology and host specificity of Spurgia capitigena (Bremi) (Dipt., Cecidomyiidae), for the biological control of Euphorbia esula L. in North America

Abstract: Spurgia capitigena (Bremi) was evaluated as a potential biological control agent of leafy spurge, Euphorbia esula L. (species complex) in North America. To ascertain the host specificity of this gall midge, tests were conducted in the field at Cavaillon, France, and at laboratories located in Rome, Italy and Bozeman, Montana, USA, in 1994 and 1995. Twenty-nine plant species, including eight native North American euphorbias and biotypes of E . esula , were evaluated. In the laboratory, the gall-midge was able to induce galls on seven species of spurges (all belonging to the subgenus Esula) and the three accessions of leafy spurge. In field tests, S . capitigena infested only E . esula . The biology and host specificity of S . capitigena from France appears to be similar to that observed for Spurgia esulae Gagné from Italy (= Bayeria capitigena Bremi). In France, this gall-midge was commonly observed from early May until October in habitats of very heavy and moist soils located along water channels, roadsides and in fruit orchards. Galls were induced on meristematic tissues, and thereby prevented flowering.     

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.     

Interaction of rhizobacteria with leafy spurge (Euphorbia esula L.) callus tissue cells

The interaction of two rhizobacterial isolates,Pseudomonas fluorescens isolate LS 102 andFlavobacterium balustinum isolate LS 105, with leafy spurge cells at the cellular level was studied using scanning and electron microscopy. Leafy spurge callus tissue inoculated with either isolate showed considerable changes compared to non-inoculated tissue. The attachment of rhizobacteria to cell surfaces was associated with the elaboration of fibrillar material which may anchor bacteria to surfaces and contribute to mediation of the phytotoxic effect caused by rhizobacteria. At the ultracellular level, inoculated callus tissue showed numerous cell alterations including esiculation and convolution of the plasmalemma, cell wall degradation and disorganization of the cytoplasm, similar to those detected in the whole plant. It is concluded that callus tissue may provide an excellent working model to investigate the mode and/or mechanism of action of potential biocontrol agents on their host plants.     

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.     

Characterization, expression and function of DORMANCY ASSOCIATED MADS-BOX genes from leafy spurge

DORMANCY ASSOCIATED MADS-BOX (DAM) genes are related to AGAMOUS-LIKE 24 and SHORT VEGETATIVE PHASE genes of arabidopsis and are differentially regulated coordinately with endodormancy induction and release in buds of several perennial plant species. DAM genes were first shown to directly impact endodormancy in peach where a deletion of a series of DAM resulted in loss of endodormancy induction. We have cloned and characterized several MADS box genes from the model perennial weed leafy spurge. Leafy spurge DAM genes are preferentially expressed in shoot tips and buds in response to cold temperatures and day length in a manner that is relative to the level of endodormancy induced by various environmental conditions. Over-expression of one DAM gene in arabidopsis delays flowering. Additionally, we show that at least one DAM gene is differentially regulated by chromatin remodeling. Comparisons of the DAM gene promoters between poplar and leafy spurge have identified several conserved sequences that may be important for their expression patterns in response to dormancy-inducing stimuli.     

Nesaecrepida infuscata: a biological control agent of the invasive plant Mimosa pigra

Mimosa pigra L. is a serious weed of wetlands of Australia, Asia and Africa. A suite of established biocontrol agents have been introduced in Australia and some Asian countries, but better control is needed. Nesaecrepida infuscata (Schaeffer) (Coleoptera: Chrysomelidae) is a common insect on M. pigra in tropical America. The larvae develop on the roots while the adults feed on the leaves. As both roots and leaves of M. pigra are relatively undamaged in the introduced range, this species has potential to limit the growth, survival and seed production. Furthermore, it is abundant in the dry season and so inflicts damage when most other agents are not active. In host specificity tests, larvae did not develop on any of the 65 test plant species other than M. pigra. Adult feeding on test plant species other than M. pigra was minimal. Based on these results, this insect has been released in Australia.     

Host‐plant use by two Orthomeria (Phasmida: Aschiphasmatini) species feeding on Macaranga myrmecophytes

Myrmecophytes depend on symbiotic ants (plant‐ants) to defend against herbivores. Although these defensive mechanisms are highly effective, some herbivorous insects can use myrmecophytes as their host‐plants. The feeding habits of these phytophages on myrmecophytes and the impacts of the plant‐ants on their feeding behavior have been poorly studied. We examined two phasmid species, Orthomeria alexis and O. cuprinus, which are known to feed on Macaranga (Euphorbiaceae) myrmecophytes in a Bornean primary forest. Our observations revealed that: (i) each phasmid species relied on two closely‐related myrmecophytic Macaranga species for its host‐plants in spite of their normal plant‐ant symbioses; and (ii) there was little overlap between their host‐plant preferences. More O. cuprinus adults and nymphs were found on new leaves, which were attended by more plant‐ants than mature leaves, while most adults and nymphs of O. alexis tended to avoid new leaves. In a feeding choice experiment under ant‐excluded conditions, O. alexis adults chose a non‐host Macaranga myrmecophyte that was more intensively defended by plant‐ants and was more palatable than their usual host‐plants almost as frequently as their usual host‐plant, suggesting that the host‐plant range of O. alexis was restricted by the presence of plant‐ants on non‐host‐plants. Phasmid behavior that appeared to minimize plant‐ant attacks is described.     

Malacorhinus irregularis for biological control of Mimosa pigra: host-specificity,life cycle,and establishment in Australia

Malacorhinus irregularis Jacoby (Coleoptera: Chrysomelidae: Galerucinae: Galerucini), from Mexico is identified as a potential biological control agent for Mimosa pigra L. (Mimosaceae), a serious weed of northern Australia and Asia. The adults feed on leaves of the host, and the larvae develop on seedlings, roots, and perhaps other plant parts. The damage to the target plant is substantial, indicating that this insect could be an effective control agent. Host-specificity tests examined the suitability of seedlings and leaves for larval development, and suitability of leaves for adult feeding. In no-choice tests, no larval development occurred on any of the 81 test plant species other than M. pigra. The extent of adult feeding on the test plants was negligible in the tests using a choice-minus-control design, being less than 1% of that which occurred on M. pigra. We conclude that M. irregularis is a specialist on its host and the risk associated with its release in Australia is low. It was released in infestations of M. pigra in the Northern Territory of Australia in 2000. Establishment and abundance was monitored at one site where the number and distribution of adult beetles fluctuated widely as soil moisture conditions varied through the seasons. Adults were found for two years after release and local defoliation of plants was attributed to this species. Although only limited observations were made, adult feeding was not recorded from Neptunia major (Benth). Windler plants growing in close proximity to M. pigra, indicating specificity under field conditions.     

Performance of Bemisia tabaci (Genn.) Biotype B (Hemiptera: Aleyrodidae) on Weeds

Bemisia tabaci (Genn.) biotype B (Hemiptera: Aleyrodidae) is regarded as a pest with a large number of hosts, including crops and weeds. The performance of this whitefly on seven weeds was evaluated in order to identify the most suitable host. The following weeds that are very common in intense agricultural areas in the state of São Paulo, Brazil, were selected for this study: spurge (Euphorbia heterophylla), beggarticks (Bidens pilosa), red tasselflower (Emilia sonchifolia), small-flower galinsoga (Galinsoga parviflora), pigweed (Amaranthus viridis), black nightshade (Solanum americanum), and morning glory (Ipomoea sp.). In free-choice tests, adult preference and oviposition were greatest on spurge. In contrast, morning glory was the least attractive and least oviposited plant. In assays carried out for egg–adult development, egg viability was greater than 87% over all weeds, whereas nymph viability ranged from 74 to 97%. The developmental period from egg to adult ranged from 26.7 to 49.1 days among the hosts under study. The lowest nymph density rate was observed for beggarticks and morning glory. Cluster analysis resulted in a single group formed by spurge, indicating its superiority as a host for B. tabaci biotype B. Even though the parameters evaluated indicate that spurge is the most suitable host among the weeds, all the others allow the reproduction of B. tabaci biotype B. For this reason, they should be observed during cropping and the intercrop period in areas infested by this whitefly.