Impact of the Arundo scale Rhizaspidiotus donacis (Hemiptera: Diaspididae) on the weight of Arundo donax (Poaceae: Arundinoideae) rhizomes in Languedoc southern France and Mediterranean Spain

Arundo donax L. (Poaceae) is native to Mediterranean Europe and invasive in the Rio Grande Basin of North America. Rhizomes from nine sites in France and Spain infested with a candidate control agent, the armoured scale Rhizaspidiotus donacis (Hemiptera: Diaspididae) weighed 50% less than those from nine sites without scale.     

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 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.     

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.     

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.     

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 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.     

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.     

The effect of Gratiana boliviana (Coleoptera: Chrysomelidae) herbivory on growth and population density of tropical soda apple (Solanum viarum) in Florida

The effect of herbivory by Gratiana boliviana Spaeth (Coleoptera: Chrysomelidae) on the invasive, tropical soda apple (TSA) (Solanum viarum Dunal, Solanaceae), was investigated using exclusion methods and by monitoring the density of G. boliviana and the weed at four locations over a period of 40 months. TSA plants protected by insecticide were taller, wider, and had greater canopy cover that unprotected plants, and plants in closed cages were taller and wider than those in open cages. Survival of plants was higher in plots protected with insecticide than in unprotected plots in both years of a 2-year study. In the population dynamics study, the initial density of TSA was 4–5 times higher at one of the locations than at the other three sites, but within 3 years, TSA density at the high density site had declined by 90%. At the three sites which initially had a low abundance of TSA, density remained low throughout the study. The intrinsic rate of increase of G. boliviana varied between –3.9 and 4.5, but over the 3-year study, was not different from zero, indicating a stable population. The intrinsic rate of increase was lower than zero for the period from October to January, and greater than zero during the January to April period. In the periods from April to July and July to October, the rate of increase was not different from zero. The implications of these results for biological control of TSA in Florida are discussed.     

Post-establishment assessment of host plant specificity of Arytainilla spartiophila (Hemiptera: Psyllidae), an adventive biological control agent of Scotch broom,Cytisus scoparius

Scotch broom, Cytisus scoparius (Fabaceae), is a shrub native to Europe that is invasive in the USA, New Zealand and Australia. The psyllid Arytainilla spartiophila has been purposely introduced to Australia and New Zealand as a biological control agent of C. scoparius, but is an accidental introduction to California. Lupines (Lupinus spp.) are the closest native taxon to Cytisus in North America, and are therefore considered to be at the highest risk for non-target damage. However, because no lupines are native to Australia or New Zealand, only one imported forage species was evaluated during prior host specificity testing. We conducted a laboratory nymphal transfer experiment, a field choice experiment and a field survey to assess risk to three lupine species (Lupinus albifrons, Lupinus bicolor and Lupinus formosus). In the laboratory, 20% of third-instar nymphs were able to develop to adulthood on L. formosus but not on the other lupine species, while 40% completed development on C. scoparius. In the field experiment, potted lupine and C. scoparius plants were placed beside large infested C. scoparius plants; oviposition occurred on all the potted C. scoparius plants, but on none of the lupines. In the field survey, no A. spartiophila eggs or nymphs were found on naturally occurring lupines growing adjacent to infested C. scoparius. The results indicate that A. spartiophila is not likely to damage or reproduce on lupines in the field. This study provides an example of how field studies can help clarify the host specificity of biological control agents.     

Biological control of Ziziphus mauritiana (Rhamnaceae): feasibility,prospective agents and research gaps

The tropical fruit tree, Ziziphus mauritiana (Rhamnaceae), a native of the Indian subcontinent, is a pasture and environmental weed in northern Australia and Fiji. In their native range, Ziziphus spp., including commercially cultivated Z. mauritiana and Z. jujuba, are subjected to a wide range of pests and diseases. The feasibility of classical biological control of this weed has not been explored to date. Effective biological control could reduce plant vigour and seed output, thereby limiting the spread of Z. mauritiana in Australia. Two Ziziphus species are native to Australia, hence, any prospective biological control agent should be specific to Z. mauritiana. Opportunistic field surveys and literature searches identified 133 species of phytophagous insects, 9 species of phytophagous mites and 12 plant pathogens on Ziziphus spp. Host records suggest the following are possibly specific to Z. mauritiana and hence are prospective biological control agents in Australia: the seed‐feeding weevil Aubeus himalayanus; the leaf‐feeding gracillariid moth Phyllonorycter iochrysis; the leaf‐mining chrysomelid beetle Platypria erinaceus; the leaf‐folding crambid moth Synclera univocalis; the leaf‐galling midge Phyllodiplosis jujubae; and the gall‐mites Aceria cernuus and Larvacarus transitans. Host range of the rust Phakopsora zizyphi‐vulgaris includes many Ziziphus species, including the native Z. oenoplia and hence would not be a suitable biological control agent in Australia. The powdery mildew Pseudoidium ziziphi, with a host range restricted to Ziziphus species, has not been reported on Z. oenoplia. All available information on the pests and diseases of Z. mauritiana are from cultivated varieties. Hence, future surveys should focus on wild Z. mauritiana in the Indian subcontinent in areas that are climatically similar to the regions of northern Australia, where it is currently most abundant.     

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.     

The effect of plant growth-promoting rhizobacteria on the growth,physiology, and Cd uptake of Arundo donax L.

In this study, plant growth-promoting potential isolates from rhizosphere of 10 weed species grown in heavy metal-contaminated areas were identified and their effect on growth, antioxidant enzymes, and cadmium (Cd) uptake in Arundo donax L. was explored. Plant growth-promoting traits of isolates were also analyzed. These isolates were found to produce siderophores and enzymes such as 1-aminocyclopropane-1-carboxylate (ACC) deaminase, and aid in solubilization of mineral nutrients and modulate plant growth and development. Based on the presence of multiple plant growth-promoting traits, isolates were selected for molecular characterization and inoculation studies. Altogether, 58 isolates were obtained and 20% of them were able to tolerate Cd up to 400 ppm. The sequence analysis of the 16S rRNA genes indicates that the isolates belong to the phylum Firmicutes. Bacillus sp. along with mycorrhizae inoculation significantly improves the growth, the activity of antioxidants enzymes, and the Cd uptake in A. donax than Bacillus alone. Highly significant correlations were observed between Cd uptake, enzymatic activities, and plant growth characteristics at 1% level of significance. The synergistic interaction effect between these organisms helps to alleviate Cd effects on soil. Heavy metal-tolerant isolate along with arbuscular mycorrhizae (AM) could be used to improve the phytoremedial potential of plants.     

The leafmining Leurocephala schinusae (Lepidoptera: Gracillariidae): not suitable for the biological control of Schinus terebinthifolius (Sapindales: Anacardiaceae) in continental USA

The host range of Leurocephala schinusae Davis & Mc Kay (Lepidoptera: Gracillariidae) was studied to assess its suitability as a biological control agent of Schinus terebinthifolius Raddi (Anacardiaceae), a serious environmental weed in the USA and elsewhere in the world. The host range was determined in the laboratory with adult no-choice oviposition (Argentina and USA) and larval development tests (USA). Seventeen plant species in ten genera were selected based on taxonomic relatedness to S. terebinthifolius, economic importance, and availability. Additional information was obtained by sampling foliage of S. terebinthifolius and six other South American native Anacardiaceae species in north-eastern Argentina. In the laboratory, except for Lithrea molleoides and Spondias mombin, all of the tested species were accepted for oviposition with a marked preference for Rhus aromatica. Incipient mines successfully developed into complete mines, pupae and adults on R. aromatica, Rhus copallinum, Schinus molle, Schinus lentiscifolius and S. terebinthifolius. In the field, although L. schinusae showed a clear preference for S. terebinthifolius, the host range, as determined by samples of host use in the native range, included three other Schinus species (S. lentiscifolius, Schinus longifolius, Schinus weinmannifolius) and one Astronium species (Astronium balansae). In conclusion, L. schinusae will not be considered for the biological control of S. terebinthifolius in continental US. However, the utilisation of this species in other infested areas such as Hawaii and Australia should be further discussed.     

The unusual cover of the armoured scale Cryptaspidiotus barbusano (Lindinger) (Hemiptera: Diaspididae: Aspidiotinae) with comments on the scale covers of other Diaspididae

Cryptaspidiotus barbusano (Lindinger 1908) is an endemic species of the Canary Islands, only known on Apollonias barbujana (Lauraceae), a plant growing in the “Laurisilva” or laurel forest, in several Macaronesian islands. Recently C. barbusano was collected on Heberdenia excelsa (Myrsinaceae), a new host plant and family, in the Jardín Botánico Canario Viera y Clavijo, Las Palmas, Gran Canaria. The recent collection of C. barbusano gives us the opportunity to redescribe its unusual adult female scale cover. The truncate cone-shaped scale cover and its peculiar reflexed exit tunnel for crawlers, with microscopic details of this rare species, are described and illustrated. The shape of the exit tunnel is discussed and the scale cover is compared with scale covers of several other diaspidid species. Finally, this study shows that C. barbusano is not a pupillarial species as formerly described.     

Laboratory host range of Austromusotima camptozonale (Lepidoptera: Crambidae), a potential biological control agent of Old World climbing fern,Lygodium microphyllum (Lygodiaceae)

Old World climbing fern, Lygodium microphyllum, is one of the most serious invasive weeds impacting south Florida and development of biological control is crucial for sustainable management. Larvae of a small moth, Austromusotima camptozonale, were discovered defoliating L. microphyllum in Australia. Preliminary testing suggested this moth was a Lygodium specialist. Laboratory host range testing was conducted on 65 species of test plants, from 31 families, comprising seven Lygodium species, four close relatives, 45 other species of ferns and fern allies, eight agricultural crops and one gymnosperm species plus the primary host L. microphyllum. Significant oviposition occurred only on other species of Lygodium. No eggs were laid on the agricultural crops, or about half the species of non-Lygodium ferns and fern allies tested. Oviposition on the other non-Lygodium ferns was very low, except on Anemia adiantifolia and Blechnum serrulatum, which received modest egg loads, but did not support development to adult. Larval feeding was low to non-existent on all the non-Lygodium species. Larvae developed to adult only on the native, American climbing fern L. palmatum, and to a lesser extent on L. japonicum. Lygodium japonicum is a naturalized invasive weed in the United States. Colonies of A. camptozonale were unable to persist on L. palmatum and died out in two to seven generations. Freezing winter temperatures in states where L. palmatum occurs would be lethal to A. camptozonale. It was concluded that A. camptozonale would pose no threat to native or cultivated plants in North America or the Caribbean and should be considered as a weed biological control agent against L. microphyllum.     

Response of the invasive pompom weed,Campuloclinium macrocephalum (Asteraceae), to feeding by the foliage-deforming thrips Liothrips tractabilis (Phlaeothripidae) under outdoor conditions in South Africa

The foliage-deforming thrips Liothrips tractabilis Mound and Pereyra (Thysanoptera: Phlaeothripidae) was recently released in South Africa as the first classical biological control agent against the invasive South American herb, Campuloclinium macrocephalum (Less.) DC. (Asteraceae). The impact of the thrips on young plants and regrowth of C. macrocephalum was assessed under outdoor conditions, and the results were compared to those of earlier laboratory trials that were conducted while the agent was still under investigation in quarantine. The outdoor trials revealed that feeding by L. tractabilis reduced the growth and biomass accumulation of C. macrocephalum, particularly in young plants and to a lesser extent in regrowth, which was largely consistent with the results of the earlier laboratory trials. Thrips-infested young plants were significantly shorter, produced fewer leaves and displayed significantly lower wet root masses and lower below- and above-ground dry masses, compared to the control plants. Although thrips-infested regrowth were significantly shorter, produced fewer leaves and displayed significantly lower above-ground dry masses than the control plants, the differences in wet root masses and below-ground dry masses were not significant. Although laboratory-based impact assessments are not necessarily an accurate reflection of field impact, in this instance the results were largely validated by the outdoor assessments. The results suggest that L. tractabilis will have a negative impact on C. macrocephalum populations in the field and thus contribute towards the weed’s biological control. The establishment of L. tractabilis has recently been confirmed in South Africa, providing the opportunity to verify these predictions.     

A new species of armored scale, Mycetaspis ailynaomi (Hemiptera, Diaspididae, Aspidiotinae), associated with Mammea americana L. (Malpighiales, Calophyllaceae) from Puerto Rico

A new species of armored scale, Mycetaspis ailynaomi Dones and Evans is described and illustrated from specimens collected on mamey (Mammea americana) from Puerto Rico. A key to the species of Mycetaspis is provided.