The effect of the armored scale,Rhizaspidiotus donacis (Hemiptera: Diaspididae), on shoot growth of the invasive plant Arundo donax (Poaceae: Arundinoideae)

The effect of feeding by the armored scale, Rhizaspidiotus donacis (Leonardi, 1920) (Hemiptera: Diaspididae) on the growth of the plant Arundo donax L. (Poaceae) was evaluated under field conditions in its native range. The study was designed to evaluate the impact of R. donacis, a candidate agent for biological control of A. donax which is invasive in arid riparian ecosystems of the Southwestern USA and Mexico. The study was carried out at five A. donax sites in the Province of Alicante, Spain, differing in altitude and climate. At each site, 30 infested lateral shoots were selected and 15 were randomly treated monthly with imidacloprid insecticide. Shoot lengths were measured monthly over a 1-year period in a comparative growth analysis. Shoots infested with R. donacis had an over 2-fold reduced growth rate as compared to treated shoots. Growth of shoots varied by site, and the effect of R. donacis on growth was most pronounced in the late spring, when mature females produced first instar scale crawlers. The impact of R. donacis on A. donax growth under field conditions in the native range, combined with its narrow host specificity, indicate that R. donacis is a promising candidate for biological control of A. donax in North America and other areas invaded by this weed.     

Evaluation of Spanish Arundo scale Rhizaspidiotus donacis (Hemiptera; Diaspididae) survival and fecundity on three new world genotypes of Arundo donax (Poaceae; Arundinoideae)

A pre-release evaluation of survival and fecundity of the arundo scale, Rhizaspidiotus donacis, was conducted on three invasive genotypes of the riparian weed, Arundo donax. The three A. donax genotypes were collected from Laredo, Austin and Balmorhea, TX, which represented the majority of the genotypic diversity found in Texas watersheds. Although R. donacis developed on all three genotypes of the plant, the Austin A. donax genotype, followed by the Laredo genotype, were the most suitable in terms of the size of first-generation immature and adult scale populations that developed after crawler release. Both the Laredo and Austin genotypes of A. donax are likely to be of Spanish origin and are close genetic matches with scale's original host plant genotype in Alicante, Spain. In comparison, survival was lowest on the phylogenetically distant genotype of A. donax from Balmorhea, TX. Although the population size of settled, immature second-generation scales varied in a manner similar to that of the first generation, the fecundity of isolated first-generation females was not significantly different across the three plant genotypes, suggesting that R. donacis is not a genotype specialist in terms of nutrient assimilation for reproduction. Rather, differences in genotype suitability affect rates of success of crawler settling. These results indicate that selection of scale genotype from the native range may have a moderate influence on the success of R. donacis and ultimately the biological control programme.     

Host range of the European,rhizome-stem feeding scale Rhizaspidiotus donacis (Hemiptera: Diaspididae), a candidate biological control agent for giant reed,Arundo donax (Poales: Poaceae) in North America

The armored scale Rhizaspidiotus donacis (Leornardi) was evaluated as a potential biological control agent of the invasive reed grass Arundo donax in North America. No-choice tests, native range field surveys and non-target host exposures were used to determine the fundamental host range of the scale collected from Caloma, Spain and Perpignan, France. Thirty-five species, including two genotypes of A. donax and seven ecotypes of Phragmites australis, along with closely related grasses, economic grasses and habitat associates were tested. In quarantine no-choice testing using releases of 200 crawlers per plant, normal development of R. donacis was observed on A. donax and A. formosana, with very limited survival to the adult stage on Spartina alterniflora and Leptochloa spp. In follow-up studies using 1000 crawlers per plant, 10 live adult females were found on Leptochloa virgata, and one adult female on Spartina alterniflora, but average adult female abundance per plant was (2580%) 26-times lower on L. virgata and over (39,090%) 100-times lower on S. alterniflora than on A. donax. Field surveys were conducted at five locations in Spain and France at which A. donax infested with R. donacis, co-occurred with two non-target species of concern and R. donacis was only found on A. donax. Six-month field host exposures in Spain using potted Leptochloa plants entwined with heavily infested A. donax confirmed that R. donacis is specific to Arundo under field conditions. Based on our results, the scale R. donacis appears to be specific to the genus Arundo and is unlikely to harm native or cultivated plants in the Americas.     

Impact of the biological control agent Tetramesa romana (Hymenoptera: Eurytomidae) on Arundo donax (Poaceae: Arundinoideae) along the Rio Grande River in Texas

Five years post-release of the arundo gall wasp, Tetramesa romana, into the riparian habitats of the lower Rio Grande River, changes in the health of the invasive weed, Arundo donax, or giant reed, have been documented. These changes in plant attributes are fairly consistent along the study area of 558 river miles between Del Rio and Brownsville, TX, and support the hypothesis that the arundo wasp has had a significant impact as a biological control agent. Plant attributes were measured prior to release in 10 quadrats at each of 10 field sites in 2007, and measured again at the same undisturbed sites, 5 years after the release of T. romana, in 2014. Above ground biomass of A. donax decreased on average by 22% across the 10 sites. This decline in biomass was negatively correlated to increased total numbers of T. romana exit holes in main and lateral shoots per site in 2014 compared to 2007. Changes in biomass, live shoot density and shoot lengths, especially the positive effect of galling on main and lateral shoot mortality, appear to be leading to a consistent decline of A. donax. Economically, this reduction in A. donax biomass is estimated to be saving 4.4 million dollars per year in agricultural water. Additional impacts are expected as populations of the wasp increase and as other biological control agents such as the arundo scale, Rhizaspidiotus donacis, become more widespread.     

Effect of water deficit on generation time and reproduction of the gall wasp Tetramesa romana,a biological control agent of giant reed (Arundo donax)

Water deficit stress can reduce the reproductive performance of galling insects, but has not previously been studied in the context of mass-rearing of a galling agent on a perennial grass. The effects of water deficit were examined for the wasp Tetramesa romana Walker (Hymenoptera: Eurytomidae), released in the Lower Rio Grande Basin of Texas and Mexico in 2009 and in northern California in 2010 for control of giant reed (Arundo donax (L)) (Poaceae). In one study, water deficit was imposed for 7–10 weeks during gall maturation, ending when adult progeny began to emerge. Aboveground water content was reduced by 1.7% and culm height by 41% in pots receiving one-fourth of normal watering (soil saturation), indicating that water deficit stress occurred. Water deficit did not affect proportion of culms successfully galled or number of progeny produced. However, time to first exit hole appearance was 2–4 days longer and adult wasp generation time 5–7 days longer on galls on water deficit–stressed compared to control culms, thereby reducing the wasp's intrinsic rate of increase. Water deficit imposed only during parent wasp oviposition had no effect on galling success or reproduction. Water deficit thus has a negative effect on rearing by delaying adult emergence and reducing the rate of population increase, even without affecting adult fertility. Mass-rearing should involve well-watered plants, and variable drought conditions in the field should be considered when evaluating T. romana establishment and impact.     

Impact of the arundo wasp,Tetramesa romana (Hymenoptera: Eurytomidae), on biomass of the invasive weed,Arundo donax (Poaceae: Arundinoideae), and on revegetation of riparian habitat along the Rio Grande in Texas

An invasive grass, Arundo donax, occupies thousands of hectares of arid riparian habitat along the Rio Grande in Texas and Mexico, and has negative impacts on national security, water resources, and riparian ecosystems. The shoot-tip-galling wasp Tetramesa romana was released in 2009 between Brownsville and Del Rio, Texas, and has dispersed over 800?km along the river channel. Plots along the river were surveyed for shoot counts of arundo and all other plant species in 2016 at seven sites in regions in which prior studies had documented a 22% decline in arundo biomass (estimated from live shoot length) from 2007 to 2014. Estimated live biomass declined a further 32% between 2014 and 2016. Native plants accounted for 86% of the 44 species encountered in plots. Individual plots averaged five plant species, and arundo was most abundant in only 9 of 21 plots. Arundo live biomass and shoot density were negatively associated with plant diversity, indicating that live arundo interferes with germination and/or survival of other plant species. The proportion of dead shoots in plots, proportion of wasp-galled shoots, and density of exit holes made by emerging adult wasps per metre live main shoot length were positively associated with plant diversity in a combined model. Regressions indicated that the effects of wasp damage measures on diversity were mediated through their effects on main shoot mortality. By reducing live arundo biomass, the arundo wasp is fostering recovery of native plant communities at riparian sites along the Rio Grande.     

Origin of the invasive Arundo donax (Poaceae): a trans-Asian expedition in herbaria

Background and Aims

The hypothesis of an ancient introduction, i.e. archaeophyte origin, is one of the most challenging questions in phylogeography. Arundo donax (Poaceae) is currently considered to be one of the worst invasive species globally, but it has also been widely utilzed by man across Eurasia for millennia. Despite a lack of phylogenetic data, recent literature has often speculated on its introduction to the Mediterranean region.

Methods

This study tests the hypothesis of its ancient introduction from Asia to the Mediterranean by using plastid DNA sequencing and morphometric analysis on 127 herbarium specimens collected across sub-tropical Eurasia. In addition, a bioclimatic species distribution model calibrated on 1221 Mediterranean localities was used to identify similar ecological niches in Asia.

Key Results

Despite analysis of several plastid DNA hypervariable sites and the identification of 13 haplotypes, A. donax was represented by a single haplotype from the Mediterranean to the Middle East. This haplotype is shared with invasive samples worldwide, and its nearest phylogenetic relatives are located in the Middle East. Morphometric data characterized this invasive clone by a robust morphotype distinguishable from all other Asian samples. The ecological niche modelling designated the southern Caspian Sea, southern Iran and the Indus Valley as the most suitable regions of origin in Asia for the invasive clone of A. donax.

Conclusions

Using an integrative approach, an ancient dispersion of this robust, polyploid and non-fruiting clone is hypothesized from the Middle East to the west, leading to its invasion throughout the Mediterranean Basin.     

Effect of nitrogen fertilization on growth of Arundo donax and on rearing of a biological control agent,the shoot gall-forming wasp Tetramesa romana

The shoot tip-galling wasp Tetramesa romana Walker (Hymenoptera: Eurytomidae) has been released for biological control of giant reed or arundo (Arundo donax L.) (Poaceae), an invasive grass in the USA and Mexico. The role of urea fertilization to improve plant-based mass-rearing was examined. In a greenhouse study, rhizomes were fertilized with urea pellets at rates equivalent to 1000 kg (low), 2000 kg (moderate), and 4000 (high) kg N per ha^–1. Total nitrogen content of ungalled stems was significantly 0.60–0.65% higher under low and moderate fertilization compared to unfertilized pots, and shoot water content was elevated 3–4% at all urea levels. Moderate fertilization significantly (by 1.4-fold) increased the relative growth rate of all shoots in pots, but did not affect final dry biomass. Fertilization did not affect number and duration of probing events by females. The percentage of shoots colonized by wasps that were galled, progeny production per shoot and per female, and emergent wasp size were not affected. However, average generation time (adult to adult) of emergent wasps was 4–5 days shorter on shoots in pots under moderate and high urea fertilization. After a four-week wasp emergence period, only 3–9% of progeny remained in fertilized shoots, while 21% of progeny remained inside unfertilized shoots. In field plots, fertilization did not affect gall density per m shoot length or per female released. Urea fertilization increased the efficiency of greenhouse rearing of the arundo wasp and availability of adults for release, even without direct effects on gall production.     

Host range of the European leaf sheath mining midge,Lasioptera donacis Coutin (Diptera: Cecidomyiidae), a biological control of giant reed,Arundo donax L.

The fundamental host range of the arundo leafminer, Lasioptera donacis a candidate agent for the invasive weed, Arundo donax was evaluated. L. donacis collects and inserts spores of a saprophytic fungus, Arthrinium arundinis, during oviposition. Larvae feed and develop in the decomposing leaf sheath channel tissue. Thirty-six closely related and economic grass species along with several key habitat associates were evaluated in no-choice tests. L. donacis and its associated saprophyte completed development only on A. donax, in concurrence with published reports from its native range in Mediterranean Europe. The arundo leafminer feeding leads to premature defoliation, constituting a different mode of attack on the host plant as compared to two previously released insects, the arundo wasp and arundo scale, which feed on shoot tips and rhizomes, respectively. Defoliation of A. donax is expected to increase light penetration into stands of A. donax which increases visibility for law enforcement, reduces the survival of cattle fever ticks, and enhance recovery of the native riparian vegetation along the Rio Grande and other habitats where this weed is invasive.     

Biology and prey range of Cryptognatha nodiceps (Coleoptera: Coccinellidae), a potential biological control agent for the coconut scale, Aspidiotus destructor (Hemiptera: Diaspididae)

Cryptognatha nodiceps Marshall is an important natural enemy used in biological control programmes against Aspidiotus destructor (Signoret), an introduced pest of coconut in many countries. In order to increase our understanding of C. nodiceps, the present study to quantify aspects of the developmental, reproductive and feeding biology under constant environmental conditions (26±2°C, 55-65% RH) was carried out. In addition, methods for culturing the scale and the beetle were developed. The average duration of development was 19.2±0.1 days for males and 19.6±0.2 days for females. Female longevity and lifetime fecundity was 35.6±5.1 days and 141.0±35.6 eggs, respectively. Life table statistics were as follows: reproductive rate, 9.99, intrinsic rate of increase, 0.09, finite rate of increase, 1.1, doubling time, 7.8 days and generation time, 41.1 days. Studies on feeding biology showed that C. nodiceps was oligophagous. The coccinellid fed and reproduced on prey species in two families, Diaspididae and Coccidae. Adults also fed on the coconut whitefly Aleurodicus cocois (Curtis) (Aleyrodidae) but no reproduction occurred on this prey. Aphididae, Psyllidae and Pseudococcidae were not fed upon.     

Host specificity assessment and potential impact of Megamelus scutellaris (Hemiptera: Delphacidae) on waterhyacinth Eichhornia crassipes (Pontederiales: Pontederiaceae)

The delphacid Megamelus scutellaris Berg was evaluated for host specificity and potential impact as part of a biological control program targeting Eichhornia. crassipes. Survival and development of adults and nymphs were used as metrics with no-choice, two-choice, nymph transfer, and sustainability tests conducted under quarantine conditions. A total of 69 plant species were tested including 12 from the Pontederiaceae (including E. crassipes). Additionally, 27 native and 5 exotic associated wetland species and 11 economic species were tested. Megamelus scutellaris exhibited a high level of oviposition and developmental fidelity to E. crassipes by failing to sustain populations on any non-target test plant past the F₁ generation. Nymph transfer tests which simulated potential spill-over events found that survival was virtually non-existent on associated wetland plants, regardless of taxonomic relatedness, including on Pontederia cordata, an important and widespread native species. Eichhornia crassipes plants exposed to two consecutive generations of feeding produced 66.9% less biomass and 73.4% fewer leaves than those in the controls. We conclude that Megamelus scutellaris is safe to release on E. crassipes in the United States.     

Comparison of two populations of Pseudophilothrips ichini (Thysanoptera: Phlaeothripidae) as candidates for biological control of the invasive weed Schinus terebinthifolia (Sapindales: Anacardiaceae)

Brazilian peppertree, Schinus terebinthifolia Raddi (Sapindales: Anacardiaceae) (hereafter Schinus), is one of the worst invasive species in Florida and Hawaii. The thrips Pseudophilothrips ichini Hood (Thysanoptera: Phlaeothripidae) is being considered as a potential biological control agent of Schinus. Two populations of this thrips were collected in the weed's native range; one from central-east Brazil (Ouro Preto thrips) and a second from north-east Brazil (Salvador thrips). Temperature requirements, adult fecundity and impact on different plant haplotypes by P. ichini were examined in the laboratory. Complete development of thrips from both populations occurred at temperatures ranging from 20 to 30°C. Two approaches were used to model the predicted distributions of the thrips populations in the USA: the physiological model (NAPPFAST) based on cold tolerance and the ecological niche model based on climatic variables (MaxEnt). The physiological model predicted that both populations of P. ichini may establish in similar areas of the USA, overlapping with the distribution of Schinus. However, the niche model predicted that only the Ouro Preto thrips could establish in the USA. The difference in model predictions suggests an apparent preadaptation of the Salvador thrips to lower temperatures than those experienced at the locations they were collected in Brazil. The Ouro Preto thrips had similar fecundity on two Florida Schinus haplotypes, whereas lower fecundity on haplotype A was found for the Salvador thrips. Based on these results, the Ouro Preto population may be better adapted to the climatic conditions and plant haplotypes found in Florida. Moreover, greenhouse studies indicated that Schinus growth was greatly reduced by thrips feeding, which may result in lower weed reproduction and densities in the field.     

Galls induced by Calophya latiforceps (Hemiptera: Calophyidae) reduce leaf performance and growth of Brazilian peppertree

Schinus terebinthifolia Raddi (Anacardiaceae) (Brazilian peppertree) is one of the most serious terrestrial invasive plants in Florida. The leaf galling psyllid, Calophya latiforceps Burckhardt (Hemiptera: Calophyidae), was discovered feeding on the Brazilian peppertree in Bahia, Brazil, in 2010, and is being evaluated for release as a biological control agent. In order to understand how this candidate biological control agent affects the growth of S. terebinthifolia, we measured the effect of the psyllid on photosynthesis, leaf chlorophyll content and plant growth. Infested plants had lower photosynthesis compared to uninfested plants 30 and 45 days after gall initiation, and chlorophyll content was lower over a 70-d period. Plant height was reduced 31% and biomass 11% after three months of infestation. Results of these studies suggest that C. latiforceps will negatively affect the growth of the Brazilian peppertree if released in Florida.     

Impact of the invasive ant Wasmannia auropunctata (Formicidae: Myrmicinae) on local ant diversity in southern Cameroon

The little fire ant, Wasmannia auropunctata Roger, is one of the world's most destructive invasive ants. It has been present in Cameroon for more than four decades, but its impact on local ant diversity is not known. We studied impact of W. auropunctata in three disturbed habitats located in rural and urban areas. We monitored ant diversity in both invaded and noninvaded zones in each area using a combination of three sampling methods: bait, pitfall traps and visual catch in quadrat. We collected 28 species in urban area and 64 in rural area. In invaded zone, W. auropunctata made up 97.72% and 99.96% of all ant fauna and ant species richness decreased to 7 and 2 in urban and rural area, respectively. In accordance with others findings in introduced environments, the presence of W. auropunctata has severely reduced abundance and richness of local ant species in both urban and rural environments in Cameroon. Measures should therefore be put in place to prevent its introduction in natural environment as forest reserves and natural parks.     

Biological parameters and thermal requirements of the parasitoid Praon volucre (Hymenoptera: Braconidae) with Macrosiphum euphorbiae (Hemiptera: Aphididae) as host

The effect of temperature on the biology of Praon volucre (Haliday, 1833) (Hymenoptera: Braconidae) in Macrosiphum euphorbiae (Thomas, 1878) (Hemiptera: Aphididae) hosts was studied and the thermal requirements of the parasitoid were determined. Experiments were carried out at 16, 19, 22, 25, and 28 ± 1°C with 70 ± 10% relative humidity and 12 h photophase. The developmental time of P. volucre males and females decreased between 16 and 23°C. The highest percentages of aphid parasitism and adult parasitoid emergence were observed at 16, 19, and 22°C. The sex ratio (expressed as % females) decreased between 16 and 23°C. Male and female longevity was high between 19 and 22°C and decreased strongly at 25°C. The lower temperature threshold for P. volucre was 5.17°C and the thermal constant was 243 degrees-days.     

Life history of Anomalococcus indicus (Hemiptera: Lecanodiaspididae), a potential biological control agent for prickly acacia Vachellia nilotica ssp. indica in Australia

Babul scale Anomalococcus indicus Ramakrishna Ayyar, a major pest of Vachellia nilotica (L.f.) P.J.H. Hurter & Mabb. on the Indian subcontinent, has been identified as a potential biocontrol agent for prickly acacia V. nilotica subsp. indica (Benth.) Kyal. & Boatwr. in Australia and was imported from southern India for detailed assessment. The life history of A. indicus under controlled glasshouse conditions was determined as a part of this assessment. Consistent with other scale species, A. indicus has a distinct sexual dimorphism which becomes apparent during the second instar. Females have three instars, developing into sexually mature nymphs after 52 days. The generation time from egg to egg was 89 days. Females are ovoviviparous, ovipositing mature eggs into a cavity underneath their body. An average of 802 ± 114 offspring were produced per female. Reproductive output was closely associated with female size; larger females produced more than 1200 offspring. Crawlers emerged from beneath the female after an indeterminate period of inactivity. They have the only life stage at which A. indicus can disperse, though the majority settle close to their parent female forming aggregative distributions. In the absence of food, most crawlers died within three days. Males took 62 days to develop through five instars. Unlike females, males underwent complete metamorphosis. Adult males were small and winged, and lived for less than a day. Parthenogenesis was not observed in females excluded from males. The life history of A. indicus allows it to complement other biological control agents already established on prickly acacia in Australia.     

Impact of the shoot-boring moth Dichrorampha odorata (Lepidoptera: Tortricidae) on growth and reproductive potential of Chromolaena odorata (Asteraceae) in the laboratory

A 9-month laboratory study was carried out to determine the impact of herbivory by a moth with shoot-boring larvae, Dichrorampha odorata Brown and Zachariades (Lepidoptera: Tortricidae) on growth and reproductive ability of its host plant, Chromolaena odorata (L.) King and Robinson (Asteraceae), a major invasive alien plant species in southern Africa. Newly hatched D. odorata larvae were inoculated onto 0 (control), 50 and 100% of the shoot tips of C. odorata in the laboratory. At all treatment levels, the basal stem diameter of C. odorata was not affected by D. odorata larval feeding. Larval feeding by D. odorata significantly reduced the height of the main shoot and flower production in C. odorata relative to the control treatment but promoted branching by increasing the number of shoot tips. However, the differences in plant height and number of flowers between the 50 and 100% inoculation levels were not statistically significant. Dichrorampha odorata is the first shoot-tip attacking agent that is being released as a biological control agent against C. odorata in South Africa. In general, the impacts of D. odorata on the weed were relatively small even though statistically significant. The findings of this study suggest that high levels of damage by the moth will modestly reduce the height, flower production, and the competitiveness of C. odorata, thereby contributing to the biological control of the weed in South Africa.     

Impact of the introduced parasitoid Peristenus digoneutis (Hymenoptera: Braconidae) on tarnished plant bug (Hemiptera: Miridae) infesting strawberries in northwestern New Jersey,USA

Lygus lineolaris populations in the northeastern USA have been markedly reduced by the introduced parasitoid Peristenus digoneutis. A 3-year study in NJ where P. digoneutis has been established for many years demonstrated that parasitism of Lygus nymphs in strawberries averaged 30% (mostly P. digoneutis). Strawberry damage by L. lineolaris ranged from 19 to 33%.