Host range and searching behaviour of Cricotopus lebetis (Diptera: Chironomidae), a tip miner of Hydrilla verticillata (Hydrocharitaceae)

A chironomid midge, Cricotopus lebetis Sublette (Diptera: Chironomidae), was discovered feeding on Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) in Crystal River, Citrus, Co., Florida, in the 1990s. Larvae of the midge mine the apical meristems of hydrilla, causing terminal branching and stunting of the plant. We investigated the fundamental host range of the midge by conducting a series of no-choice and paired-choice tests. No-choice developmental tests with neonate larvae revealed that the fundamental host range of C. lebetis included not only on hydrilla but also several other aquatic plants in different families, suggesting that this insect is not a hydrilla specialist. In paired-choice bioassays, larval colonisation of Elodea canadensis Michx. (Hydrocharitaceae) and Najas guadalupensis (Spreng.) Magnus (Najadaceae) was greater than colonisation of H. verticillata. Behavioural bioassays in a Y-tube olfactometer and in Petri dishes suggested that neonate larvae were not able to locate host plant material, whereas older larvae were successful in finding hosts. In paired-choice oviposition tests, adult females discriminated between potential oviposition sites, with greater numbers of eggs laid on E. canadensis and N. guadalupensis than on H. verticillata. This study is the first detailed account of host searching and oviposition behaviour of a phytophagous chironomid midge. The results will be used to assess the potential value of C. lebetis as a biological control agent of hydrilla.     

Effects of Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) growing conditions and nutrient content on the performance of a leaf-mining fly,Hydrellia purcelli Deeming (Diptera: Ephydridae)

Host plant quality for insects used in weed biological control influences their performance and hence their ability to suppress target host populations. Determining the specific response of these insects to the quality of their host is important because phytophagous insects have variable tolerances of the different constituents of host plant quality, most notably dietary nitrogen (N), but also other physical and chemical components. The invasive aquatic weed Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) was cultivated under varying nutrient conditions to determine the influence of plant quality on immature survival, development, larval mining, reproductive output and adult longevity of a leaf-mining fly Hydrellia purcelli Deeming (Diptera: Ephydridae). Additionally, field-collected H. verticillata was included in the investigations to assess the potential performance of H. purcelli in the field. Variation in plant tissue N and phosphorus (P) concentrations had no effects on larval survival, female fecundity, or adult longevity, but high levels of N and P were associated with reduced immature development times and higher body mass of females. Overall, plant quality factors not measured in this study appeared to have a greater impact on the performance of the fly, rather than dietary N and P. The results provided insights into optimal mass-rearing conditions for H. purcelli and the potential performance of the fly in the field in South Africa. Furthermore, the results demonstrate the importance of considering other aspects of plant quality for insect agents, in addition to dietary N and P, when developing mass-rearing protocols or predicting their potential impact in the field.     

Biology of Polypedilum n. sp. (Diptera: Chironomidae), a promising candidate agent for the biological control of the aquatic weed Lagarosiphon major (Hydrocharitaceae) in Ireland

Lagarosiphon major (Ridl.) Moss ex Wager (Hydrocharitaceae), an aquatic macrophyte native to Southern Africa that has become invasive in several countries worldwide, is a potential target for biological control. Biology studies were conducted on Polypedilum (Pentapedilum) n. sp. near reei Oyewo & Sæther (Diptera: Chironomidae), a midge whose larvae were discovered mining the plant's shoot tips in its native range. Field surveys indicated that the midge occurred only at a small number of sites but attained high densities (up to 370 shoots damaged/m²) that prevented further growth from the shoot tips. A population of the midge was imported into quarantine in Ireland to evaluate its potential as a candidate biological control agent. The adult stage is terrestrial and short-lived (4–5 d), with females depositing one–two egg packets into the water bodies. First-instar larvae fed externally on the stems and leaflets. Later instars fed on the apical meristems and burrowed into the shoot tips, with resultant damage stunting the apical growth. Larvae moved readily between shoots to complete their development and pupated within the tunnels excavated by the late instar larvae. Developmental time to adulthood ranged from 31 to 49 days at 20.7°C and a 15 L:9-D cycle. This is the first time that a chironomid has been imported and successfully cultured for consideration as a classical biological control agent. Several aspects of the midge's biology suggest that host specificity testing is warranted to determine its potential as a biocontrol agent of L. major.     

Biology and ecology of Brazilian elodea (Egeria densa) and its specific herbivore, Hydrellia sp., in Argentina

Egeria densa (Hydrocharitaceae) is a submerged macrophyte from South America that is a weed in several countries. It crowds out native plants and hinders water use, causing economic and environmental damage. The leafminer fly Hydrellia sp. 1 (Diptera: Ephydridae), was found feeding in E. densa throughout its Argentine distribution, and is currently the only known specialist herbivore of E. densa. It was reared in the laboratory and tested on 25 plant species. This herbivore can cause heavy defoliation in the laboratory and in the field. Hydrellia sp. 1 was found only on E. densa, but in the laboratory it also developed on two other Hydrocharitaceae species in the same family; Egeria naias, and Elodea callitrichoides. Significant oviposition and feeding were only observed on its primary natural host, and to a lesser degree on E. naias. Field studies indicate Hydrellia sp. 1 is present in the field year round, unless the host plant is prostrate for long periods, or covered by floating macrophytes. These results indicate Hydrellia sp. 1 may be a suitable biocontrol candidate for E. densa.     

Competition between two invasive Hydrocharitaceae (Hydrilla verticillata (L.f.) (Royle) and Egeria densa (Planch)) as influenced by sediment fertility and season

Competition between two invasive plants of similar growth form, Hydrilla verticillata (L.f.) (Royle) and Egeria densa (Planch), was studied in response to season and sediment fertility. These two invasive species were grown in outdoor concrete tanks in monocultures and mixtures. Five fertilization rates were tested for monocultures and two for mixtures where six combinations of planting densities were used in two seasons (spring and fall). Monitoring of plant biomass was made at the end of each of these 2-month-experiments. In contrast to E. densa, clear seasonal patterns in biomass production and in reproductive allocations of H. verticillata were evident. Competitive pressure for both species was lower during the fall experiment. Biomass production increased with fertilization for H. verticillata in monocultures and changes either in allocative ratios or in tuber production patterns were shown in response to nutrient availability. However, E. densa growth was not affected by fertilization. In most cases, H. verticillata was a better competitor than E. densa except when sediment was pure sand. Competition occurred mainly for nutrient uptake rather than for light harvesting. These results suggest that despite the similar ecology, H. verticillata may outcompete E. densa in many situations, probably due to its higher plasticity.     

Risk assessment of Episimus unguiculus (Lepidoptera: Tortricidae), a biological control agent of Schinus terebinthifolia (Sapindales: Anacardiaceae) in Hawaii,USA

Schinus terebinthifolia Raddi (Anacardiaceae) is an introduced ornamental tree from South America that has become one of the most invasive weeds in Hawaii and Florida, USA. Exploratory surveys in the plant’s native range from 1950 to 2014 identified several potential biological control agents. One of these is the leaflet rolling moth Episimus unguiculus Clarke (Lepidoptera: Tortricidae), previously known as Episimus utilis Zimmerman. This biological control agent was released in Hawaii in the 1950s where high densities were occasionally observed, leading to partial control of S. terebinthifolia by the 1960s. Larvae are leaf tiers capable of completely defoliating small plants. In order to investigate the release of E. unguiculus in the continental USA, a series of laboratory no- choice, and multiple-choice tests were conducted in Florida, and a preliminary open field test with a native plant in Hawaii. Under the confined laboratory conditions imposed during the no-choice tests, E. unguiculus accepted the economically important Pistacia spp. and several other non-target plants for oviposition and development. However, in the multiple-choice tests E. unguiculus exhibited a clear preference for S. terebinthifolia relative to non-target plants accepted in the no-choice tests. Overall, the results of field observations during surveys in South America and Hawaii and host range studies completed in Hawaii and Florida showed that E. unguiculus is a narrow specialist on S. terebinthifolia, its natural host plant.     

Assessing the risk of Eccritotarsus eichhorniae to pickerelweed,Pontederia cordata in North America

The mirid Eccritotarsus eichhorniae is a newly described species with potential as a biological control agent for water hyacinth, Eichhornia crassipes, in North America. A series of quarantine-based studies quantified E. eichhorniae usage of Pontederia cordata with no-choice, adult choice, nymph choice, multi-generation, and impact tests. Although greater numbers of E. eichhorniae adults developed on E. crassipes in two separate no-choice trials, some E. eichhorniae were able to complete development on P. cordata in both trials. Adults showed a clear oviposition preference for E. crassipes and F₁ progeny reduced the mean relative growth rate of E. crassipes but not P. cordata. Naïve nymphs were more likely to leave P. cordata for E. crassipes but not all nymphs left P. cordata when given a choice. Both multi-generation trials showed a rapid increase in the density of adults and nymphs on E. crassipes over four to five generations while population densities on P. cordata did not change, indicating that this insect can maintain populations on this plant. The modest damage inflicted by a sibling species on E. crassipes, coupled with the ability of E. eichhorniae to persist on P. cordata, may pose an interpretation challenge to U.S. regulatory agencies concerned with non-target impacts by biological control agents. Another option for the continuing efforts to increase the biological control of E. crassipes is to prioritise the evaluation of other potential agents like Taosa longula Remes Lenicov (Homoptera: Dictopharidae) or Thrypticus truncatus Bickel and Hernández (Diptera: Dolichopodidae) over E. eichhorniae.     

Suppression of the aquatic weed Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) by a leaf-cutting moth Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) in Jozini Dam,South Africa

A classical biological control programme against Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) was initiated in South Africa following the discovery of extensive mats in Jozini Dam in northern KwaZulu-Natal (KZN) in 2006. However, in mid-2008, high densities of a phytophagous insect, Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) were discovered on the H. verticillata infestation, causing high levels of defoliation and dieback of the weed. In order to quantify the impact of the moth and to assess the long-term threat of H. verticillata to South African water bodies, a monitoring programme was initiated in 2013. Although P. diminutalis did not demonstrate a preference for H. verticillata over a native aquatic plant, Potamogeton schweinfurthii A. Bennett (Potamogetonaceae), moth populations followed a cyclical pattern of abundance, with rapid population increases coinciding with increases in the abundance of H. verticillata. High damage levels to H. verticillata were associated with high densities of immature P. diminutalis, which consistently led to population crashes of H. verticillata. Parapoynx diminutalis appears to have great potential to contribute to the management of H. verticillata in South Africa. However, the moth's oligophagous habits will probably preclude its intentional introduction into water bodies with H. verticillata where it does not already occur.     

Susceptibility of neonate Lissachatina fulica (Achatinidae: Mollusca) to a US strain of the nematode Phasmarhabditis hermaphrodita (Rhabditidae: Nematoda)

ABSTRACT

The nematode Phasmarhabditis hermaphrodita is a lethal parasite of many pest slug and snail species. It is used as a commercial biological control agent (Nemaslug®) in Europe but a number of key gastropod pests (e.g. Lissachatina fulica) are thought to be unsusceptible. Our data, however, demonstrated that a recently discovered U.S. strain of P. hermaphrodita causes statistically significant mortality to neonate L. fulica and consequently this nematode could be an important IPM tool for managing this snail pest in areas where P. hermaphrodita has been discovered. Soil moisture content appears to play an important role in determining nematode efficacy.     

Patterns of aquatic weed regrowth following mechanical harvesting in New Zealand hydro-lakes

Mechanical harvesting is used to control submerged aquatic weeds in parts of the hydro-lakes in New Zealand's North Island. Problem species are Egeria densa and Lagarosiphon major (Hydrocharitaceae), and Ceratophyllum demersum. Experiments were conducted in two contrasting hydro-lakes. Lake Aratiatia; clear water (K _o 0.2 m^–1) and a low residence time (< 8 h), and Lake Ohakuri; turbid water (K _o = 0.6) and a longer residence time (> 5 days). Growth rates were measured underwater in harvested and control (unharvested) plots. Regrowth of C. demersum was dependent on the prior establishment of the rooted Hydrocharitaceae. Regrowth of the Hydrocharitaceae was inhibited where significant water movement occurred. Regrowth declined after 3 six-monthly harvests allowing the establishment of low growing native Nitella spp. beds in the smaller clear water lake. In Lake Ohakuri there was a change in species dominance from Ceratophyllum to Elodea canadensis in shallow (1–2 m) water. No change in species dominance was observed in deeper (>2 m) water and native species were not able to re-establish. The recommended cutting frequency for management of surface weed growths was only once per year in Lake Aratiatia, but twice per year in Lake Ohakuri.     

Natural enemies from South Africa for biological control of <Emphasis Type="Italic">Lagarosiphon major</Emphasis> (Ridl.) Moss ex Wager (Hydrocharitaceae) in Europe

The non-native invasive plant, Lagarosiphon major (Hydrocharitaceae) is a submersed aquatic macrophyte that poses a significant threat to water bodies in Europe. Dense infestations prove difficult to manage using traditional methods. In order to initiate a biocontrol programme, a survey for natural enemies of Lagarosiphon was conducted in South Africa. Several phytophagous species were recorded for the first time, with at least three showing notable promise as candidate agents. Amongst these, a leaf-mining fly, Hydrellia sp. (Ephydridae) that occurred over a wide distribution causes significant leaf damage despite high levels of parasitism by braconid wasps. Another yet unidentified fly was recorded mining the stem of L. major. Two leaf-feeding and shoot boring weevils, cf. Bagous sp. (Curculionidae) were recorded damaging the shoot tips and stunting the growth of the stem. Several leaf-feeding lepidopteran species (Nymphulinae) were frequently recorded, but are expected to feed on a wide range of plant species and are not considered for importation before other candidates are assessed. The discovery of several natural enemies in the country of origin improves the biological control prospects of L. major in Europe.     

The problematic systematic position of Ceratostratiotes Gregor (Hydrocharitaceae ?) – morphological, anatomical and biochemical comparison with Stratiotes L.

 The enigmatic fossil taxon Ceratostratiotes sinjanus (Kerner) Bužek has been considered as either a Hydrocharitaceae seed (monocot) or a Ceratophyllaceae fruit (dicot). The co-occurrence of seeds of Stratiotes kaltennordheimensis (Zenker) Keilhack (Hydrocharitaceae) and Ceratostratiotes in Early Miocene sediments in Langau (Lower Austria) enabled comparisons of morphological-anatomical features and lignin compositions, independent of diagenetic biases. The biochemistry of the Ceratostratiotes seed coat wall is not monocotyledon-like and is dissimilar to that of the co-occurring Stratiotes testae. In contrast, the Ceratostratiotes seed coat anatomy resembles that of Hydrocharitaceae genera with the micropyle and raphe being very similar to those in seeds of Stratiotes. Although the horizontal arrangement of the Ceratostratiotes spines is different from the longitudinal arrangement of the surface ridges in Stratiotes kaltennordheimensis and the spines in Blyxa (Hydrocharitaceae), the spine structures are very similar. A mixture of similarities and differences characterizes the morphological-anatomical features. Thus the systematic affinity of Ceratostratiotes still remains enigmatic. Received April 18, 2002; accepted October 18, 2002 Published online: February 4, 2003     

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

ABSTRACT

Prickly acacia, Vachellia nilotica ssp. indica (Benth.) Kyal. & Boatwr, is a significant weed of northern Australia and has been a target of weed biological control in Australia since the 1980s. Following native range surveys in India, the scale insect Anomalococcus indicus Ayyar was identified as the most promising agent and was imported into Australia for further research. A. indicus is a major pest of prickly acacia on the Indian subcontinent, where it causes shoot tip dieback and plant death. Despite field observations suggesting the species was specific to V. nilotica, A. indicus completed development on 17 of the 84 non-target plant species tested during no-choice host specificity trials under quarantine conditions. Of these, Acacia falcata, V. bidwillii, V. sutherlandii and Neptunia major supported high numbers of mature females in all replicates. All of these species were utilised in choice trials. Combined risk scores indicate that V. sutherlandii, N. major and A. falcata may be attacked in the field. Due to the limited ability of scale insects to disperse, only those non-target species that occur on the Mitchell grass downs (i.e. V. sutherlandii) are considered to be at risk. Nevertheless, in view of the disparity between quarantine test results and the observed field host specificity of A. indicus in India, field trials are currently being conducted in India using Australian native species on which complete development has occurred. The future of A. indicus as a biological control agent for prickly acacia in Australia will be determined once results from these field trials are known.     

Host range tests reveal Paectes longiformis is not a suitable biological control agent for the invasive plant Schinus terebinthifolia

The most critical step during a weed biological control program is determination of a candidate agent’s host range. Despite rigorous protocols and extensive testing, there are still concerns over potential non-target effects following field releases. With the objective to improve risk assessment in biological control, no-choice and choice testing followed by a multiple generation study were conducted on the leaf-defoliator, Paectes longiformis Pogue (Lepidoptera: Euteliidae). This moth is being investigated as a biological control agent of Schinus terebinthifolia Raddi (Sapindales: Anacardiaceae), which is one of the worst invasive plant species in Florida, USA. Results from no-choice testing showed higher larval survival on S. terebinthifolia (48 %) and its close relative Schinus molle L. (47 %), whereas lower survival was obtained on six non-target species (<25 %). When given a choice, P. longiformis females preferred to lay eggs on the target weed, but oviposition also occurred on four non-target species. An improved performance on the native Rhus aromatica Aiton was found when insects were reared exclusively on this non-target species for one or two generations. Results from host range testing suggest that this moth is oligophagous, but has a preference for the target weed. Non-target effects found during multiple generation studies indicate that P. longiformis should not be considered as a biological control agent of S. terebinthifolia.     

Invasion of the gall mite Aceria genistae (Acari: Eriophyidae), a natural enemy of the invasive weed Cytisus scoparius,into California,U.S.A. and predictions for climate suitability in other regions using ecological niche modelling

Scotch broom (Cytisus scoparius (L.) Link) is a European shrub that has naturalised in several countries worldwide and is recognised as an invasive weed in much of western North America. The mite Aceria genistae (Nalepa) is a coevolved, gall-inducing herbivore associated with Scotch broom in its native range and has been intentionally introduced as a classical weed biological control agent of C. scoparius in Australia and New Zealand. An adventive, never intentionally introduced, population of A. genistae was discovered in Washington and Oregon, U.S.A. in 2005. Surveys for A. genistae in California resulted in the discovery of the gall mite in 11 counties, with a widely scattered distribution. Molecular and morphological assessments confirm the mites collected from galls in California are A. genistae. Whether natural or anthropogenic, the estimated rate of long range dispersal for A. genistae from Washington or Oregon to California ranges from 39 to 62?km/yr. Niche model predictions indicate that A. genistae will continue to expand its distribution throughout much of the Scotch broom-invaded lands of California but areas supporting the weed in the Eastern U.S.A. appear less suitable. Modelling evidence also indicates that portions of Chile and Argentina are suitable for colonisation by A. genistae, also suggesting that expansion of the mite is possible in areas of Tasmania, southeastern Australia, and New Zealand where the mite was released. The environmental safety of A. genistae in relation to non-target plants and the influence of herbivory on Scotch broom fitness are discussed.     

Limnological effects of Egeria najas Planchon (Hydrocharitaceae) in the arms of Itaipu Reservoir (Brazil, Paraguay)

The limnological effects resulting from the metabolism of Egeria najas Planchon (Hydrocharitaceae) were evaluated in the arms of the Itaipu Reservoir (Brazil). Due to local population decline in biomass and coverage, decomposition was the main process that accounted for the observed limnological changes. Thus, total Kjeldahl nitrogen and total phosphorus in littoral regions were significantly higher during periods of E. najas occurrence. After plant stands disappeared, concentrations decreased in all arms. The nutrient pump effect explains these results, and E. najas stands probably act as sources of nutrients, especially nitrogen. However, low concentrations measured after aboveground biomass disappearance indicate that the internal loading effect is ephemeral and restricted to the littoral regions subjected to the action of naturally senescing populations. The underwater light regime was negatively affected by E. najas occurrence. Overall, these results and the small areas colonized by E. najas in the arms of the Itaipu Reservoir indicate that ecological problems (e.g., reduction of biodiversity) derived from E. najas metabolism are unlikely.     

Review of the biology and ecology of a ragweed leaf beetle,Ophraella communa (Coleoptera: Chrysomelidae), which is a biological control agent of an invasive common ragweed,Ambrosia artemisiifolia (Asterales: Asteraceae)

A ragweed leaf beetle, Ophraella communa (Coleoptera: Chrysomelidae), has been highlighted as a potential biological control agent of Ambrosia artemisiifolia. O. communa and A. artemisiifolia are native in North America and alien species in East Asia and Europe. As an invasive weed, A. artemisiifolia causes severe economic losses as reducing agricultural production as well as producing severe allergenic pollen. As an herbivore insect, O. communa has strong host preference on A. artemisiifolia. All the developmental stages of O. communa can be found on A. artemisiifolia and it attacks a single plant in repeated and extended manners. With few individuals on A. artemisiifolia, O. communa can completely defoliate before pollen production. Therefore, O. communa had been focused as a biological control of this invasive weed, but its introduction was denied because of possible damage on an important crop, Helianthus annuus. O. communa was accidentally introduced in East Asia and Europe in 1990s and 2010s, respectively. Fortunately, O. communa population was well established to suppress A. artemisiifolia in the introduced areas. Following detailed field surveys and host specificity tests of O. communa were conducted and proved a strong potential of O. communa as a biological control agent of A. artemisiifolia. Moreover, O. communa has been investigated in physiological and evolutionary studies. In this study, the potential of O. communa as a biological control agent and a study organism are reviewed.     

Life table of Gaeolaelaps aculeifer (Acari: Laelapidae) feeding on larvae of Lycoriella auripila (Diptera: Sciaridae) with stage-specific estimates of consumption

Gaeolaelaps aculeifer (Canestrini, 1883) is a soil-dwelling predatory mite with potential for use as a biological control agent of fungus gnats (Diptera: Sciaridae) in mushroom production. The life table, predation rate and population growth rate of G. aculeifer on a diet of larvae of the sciarid fly, Lycoriella auripila, at 23?±?1°C, 60?±?5% RH and a photoperiod of 0:24 (L:D)?h was investigated. The results revealed that the duration of egg, larva, protonymph, deutonymph, females and males of G. aculeifer were 3.8?±?0.1, 1.4?±?0.1, 3.9?±?0.1, 4.1?±?0.1, 67.7?±?2.8 and 60.3?±?3.1 days, respectively. Net reproductive rate (R₀) was 54.8?±?7.1 offspring, intrinsic rate of increase (r) was 0.12?±?0.01 offspring day^?1, finite rate of increase (λ) was 1.13?±?0.01 day^?1and mean generation time (T) was 32.3?±?0.6 days. The predator consumed a mean of 0.08?±?0.05, 1.73?±?0.18, 3.16?±?0.28 and 75.9?±?7.1 third instar L. auripila larvae during the larval (1.3?±?0.1 days), protonymph (3.9?±?0.1 days), deutonymph (4.1?±?0.1 days) and adult (52.6?±?2.2 days) stages. Population parameters and consumption rates suggest that G. aculeifer has good potential as a biological control agent of L. auripila in mushroom production.     

Hellcotytencnus spp. (Nematoda: Tylenchida) from managed soils in New Zealand

Abstract

Helicotylenchus species collected from managed soils in New Zealand were identified. They were found at 159 sites. H. pseudorobustus was found at 82 of these sites, H. labiatus at 56, H. varicaudatus at 28, and H. canadensis at 4. Only 11 sites had more than one species present and most (7) of these sites had mixtures of H. pseudorobustus and H. labiatus. H. labiatus, reported only twice before, was the most common species found in cotula bowling greens. H. canadensis was restricted to sites south of 45°S. Measurements are given for 15 populations, and H. labiatus is illustrated. Characters separating the four species are given.