Assessing Eucryptorrhynchus brandti (Coleoptera: Curculionidae) as a potential carrier for Verticillium nonalfalfae (Phyllachorales) from infected Ailanthus altissima

Two potential biological control agents for the invasive tree-of-heaven, Ailanthus altissima (Mill.) Swingle, have been extensively studied: a vascular wilt fungus, Verticillium nonalfalfae Inderb. (formerly Verticillium albo-atrum Reinke and Berthold), and a host-specific weevil from China, Eucryptorrhynchus brandti Harold, which is currently pending quarantine release. In 2002, V. nonalfalfae was observed in Pennsylvania causing significant mortality to A. altissima. Quarantine experiments were conducted to test different forms of transmission of the fungus by E. brandti to A. altissima. In one experiment, 75% of adult E. brandti transmitted V. nonalfalfae to A. altissima seedlings after contact with a culture and feeding on infected plant material. In another study, 14.7±1.3% of adult E. brandti ingested and passed viable V. nonalfalfae propagules into faeces after feeding on infected A. altissima stems for 24, 48 and 72 h, respectively, and the number of propagules increased at 72 h. Eighty-three per cent of surviving weevils overwintering in naturally infested potting mix carried viable V. nonalfalfae propagules externally. All weevil progeny that emerged from infected A. altissima billets appeared to be as healthy as weevils reared from non-infected billets. Our studies have shown that ingestion of V. nonalfalfae after feeding on infected plant material plays an important role in assessing E. brandti as a potential carrier. Results from these laboratory studies indicate E. brandti has the ability to transfer V. nonalfalfae to A. altissima seedlings in a laboratory setting.     

Application of a frequency distribution method for determining instars of Eucryptorrhynchus brandti (Coleoptera: curculionidae) from several morphological variables

Eucryptorrhynchus brandti (Harold) (Coleoptera: Curculionidae) is a major pest of Ailanthus altissima (Mill.) Swingle (Simaroubaceae), commonly known as tree-of-heaven, and A. altissima var. Qiantouchun in China. It is considered a potential biological control agent for tree-of-heaven in North America. The aim of this study was to use a frequency distribution method to determine the instars of field-collected larvae of E. brandti. We collected larval samples of various sizes from both the field and the laboratory and measured five morphological variables, including antenna spacing, mandible width, head-capsule width, ocellus spacing, and pronotum width. Based on the results of a frequency distribution method and Dyar’s rule, the larvae of E. brandti were divided into seven instars. Of the five variables measured, the width of the head capsule provided the best measurement for determining instar stage. The regression equation between the head-capsule widths and the instar number was y?=?0.324e^0.096x (R²?=?0.970).     

Phototactic behaviour of Eucryptorrhynchus scrobiculatus and E. brandti (Coleoptera: Curculionidae) adults*

Eucryptorrhynchus scrobiculatus and E. brandti (Coleoptera: Curculionidae) are destructive weevils on Ailanthus altissima in China. This study examined phototactic behaviour of E. scrobiculatus and E. brandti in response to eight light-emitting diodes (LEDs) in the laboratory and field. Effects of gender, starvation, and light and dark experience on the phototactic behaviour of the insects were evaluated. The results demonstrated that, the two species of weevil were phototactic insects and most sensitive to violet light (400–405?nm), followed by blue–violet light (420–430?nm). They were less sensitive to red (655–660?nm), white (6000–6500?k), blue–green (470–480?nm), yellow (590–595?nm), blue (450–455?nm), and green (515–530?nm) light. In the light intensity range of 200–1000?lux, the light intensity had no significant effect on the phototatic behaviour of E. scrobiculatus and E. brandti. Phototactic behaviour of the insects was affected by gender. The phototaxis indices of the two species of weevil increased with starvation, reaching a plateau after 2 or 3?d of starvation. The phototaxis indices of E. scrobiculatus and E. brandti were significantly affected by various wavelengths of light following exposure to the light for 3?h or in different dark experience time. In the mark-release-recapture test, the number of E. scrobiculatus and E. brandti adults trapped by violet (400–405?nm) light traps is the largest. The information provided here provides a basis for survey and control of E. scrobiculatus and E. brandti.     

Micro-habitat niche differentiation contributing to coexistence of Eucryptorrhynchus scrobiculatus Motschulsky and Eucryptorrhynchus brandti (Harold)*

Temporal, spatial, and trophic niche differentiation usually occurs among species living in similar habitats which can reduce interspecific competition under limited resources. Eucryptorrhynchus scrobiculatus and Eucryptorrhynchus brandti are sympatric and closely related species and are host-specific to Ailanthus altissima (Mill.) Swingle in China. However, the possibility of niche differentiation between the two species is not clear. Therefore, we investigated the density of the two weevils temporally, spatially, and trophically (plant parts of A. altissima) at Lingwu farm and Haojiaqiao town, Lingwu city, Ningxia Hui Autonomous Region. No differences were found in temporal Levin’s niche breath index values ranging from 3.74 to 3.95 and temporal Pianka’s niche overlap values (0.98–0.99). There was little differentiation in the vertical spatial niche. The vertical distribution of E. scrobiculatus was mainly located from 0?cm (ground) to over 250?cm on tree-of-heaven. Eucryptorrhynchus brandti was found predominantly from 0.1 to 50?cm to over 250?cm on tree-of-heaven. The overlap of the cardinal direction spatial niche was 0.90–0.98, indicating no significant overlap in the cardinal direction at Lingwu farm and Haojiaqiao town. Significant differentiation of trophic niche occurred with E. scrobiculatus feeding on 1-year-old branches, perennial branches, and petioles, while E. brandti feed on the stem.     

Oviposition site selection in Cactoblastis cactorum (Lepidoptera): constraints and compromises

Summary Oviposition by Cactoblastis cactorum on Opuntia ficus-indica and O. aurantiaca was assessed from the positioning of egg sticks on plants in the field. The number of egg sticks laid on O. ficus-indica plants was affected by: (1) plant size; (2) moth emergence near the plant; (3) cladode condition; and (4) plant conspicuousness. These factors contributed towards the clumping of egg sticks on plants. There was no apparent oviposition preference for one of the two host plant species despite the fact that egg predation was higher and fecundity lower on O. aurantiaca. The selection of a site for oviposition on the host plants was influenced by: (1) cladode condition; (2) height above ground; and (3) shelter from wind during oviposition. Succulent cladodes were the favoured sites for oviposition. The evidence suggests that in C. cactorum, oviposition site selection is largely the net result of a compromise between oviposition behaviour selected for increasing the probability of juvenile survival and oviposition behaviour selected for increasing the probability of laying the full complement of eggs. In addition, environmental and physiological factors such as wind and wing-loading, are thought to place constraints on the range of sites available for oviposition.     

Effects of starvation on death‐feigning in adult Eucryptorrhynchus brandti (Coleoptera: Curculionidae)

In insects, death‐feigning is an effective defence strategy. Eucryptorrhynchus brandti, a major borer pest in China, has a weak flight ability and exhibits obvious death‐feigning behaviour when disturbed. Despite a large number of studies of its biological and ecological properties as well as control methods, the death‐feigning behaviour has not been specifically described. In laboratory conditions, we recorded the survival rate under starvation and feeding conditions and evaluated the effect of starvation on the duration and occurrence of death‐feigning. In a continuous experiment, we examined variation in the death‐feigning duration every day over 7 day. Then, we evaluated the effects of starvation for 3, 6 and 9 day in a non‐continuous experiment and further observed variation in the death‐feigning intensity. We found that starvation significantly affected the survival rate. Survival time was significantly longer in the starvation group than in the feeding group, and females had longer survival times than males (female: 14 day, male: 8 day). In the continuous experiment, starved E. brandti had the longest duration of death‐feigning at 2 day, followed by a significant decrease. In the non‐continuous experiment, the duration and proportion of death‐feigning decreased significantly as the duration of starvation increased and were significantly lower than feeding. These observations suggest that starvation is a non‐negligible factor in the death‐feigning behaviour of E. brandti adults, facilitating the interpretation of future ecological and behavioural data of thanatosis.     

Oviposition behaviour of the soil mite Veigaia cerva (Acari: Veigaiidae)

We observed the oviposition behaviour of the soil mite Veigaia cerva (Kramer) (Acari: Veigaiidae) using continuous video-monitoring. Five phases could be recognized. Phase I involved inspection of the substrate. In phase II the female rhythmically moved her gnathosoma and first pair of legs. After an inactive phase III, the soma was raised (IV), and the egg was laid (V). In the actual egg laying three sub-phases could be distinguished: internal egg movement, placing the egg in front of the gnathosoma, and depositing the egg using the chelicerae. The palps and first pair of legs were used to position the egg between the chelicerae. The whole process took on average 333 ± 22 s.     

Biology of Bactrocera (Zeugodacus) tau (Walker) (Diptera: Tephritidae)

The biology of the fruit fly Bactrocera tau, an important horticultural pest, was studied under laboratory conditions at 25°C and 60–70% relative humidity on Cucurbita maxima. The duration of mating averaged 408.03 ± 235.93 min. After mating, the female fly had a preoviposition period of 11.7 ± 4.49 days. The oviposition rate was 9.9 ± 8.50 eggs and fecundity was 464.6 ± 67.98 eggs/female. Eggs were elliptical, smooth and shiny white, turning darker as hatching approached, and measured 1.30 ± 0.07 mm × 0.24 ± 0.04 mm. The chorion has polygonal microsculpturing and is species-specific with polygonal walls. The egg period lasts for 1.3 ± 0.41 days. The duration of the larval period is 1.2 ± 0.42, 1.7 ± 0.48 and 4.0 ± 0.94 days for first, second and third instars, respectively. Pupation occurs in the sand or soil and pupal periods are 7.0 ± 0.47 days. The life cycle from egg to adult was completed in 14.2 ± 1.69 days; the longevity of mated females and males was 130.33 ± 14.18 and 104.66 ± 31.21 days, respectively. At least two to three generations were observed from June 2008 to June 2009.     

Insecticidal and biological efficacy of two vegetable oils against Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchidae) under laboratory conditions

Bioefficacy of two vegetable oils, namely ethyl oleate and Acorus calamus, were conducted against the pulse beetle Callosobruchus maculatus (Fabr.) under a constant temperature of 29 ± 1 °C and a relative humidity of 65 ± 5% R.H. Four concentrations (0.125, 0.25, 0.5 and 1.0%) of each oil were used against the adult and tested their latent effect on the immature stages of the beetle. The results obtained show insecticidal effect of both ethyl oleate and A. calamus oil; however, A. calamus oil proved to be more toxic to the adults at all the concentrations tested in comparison to the ethyl oleate oil. As the concentration of the oils decreased, the percentage mortality decreased. The obtained results concentrated on biological efficacy of the oils tested against the adults. Both oils affected the longevities of both male and female C. maculatus adults in comparison to the control. The oviposition periods of the adult female were affected significantly in comparison to the control. A. calamus oil prohibited oviposition completely at high concentrations of 0.5 and 1.0% and caused complete sterility to the females. Lower concentrations decreased the oviposition period greatly and reduced the number of eggs laid in comparison to the control. Ethyl oleate had an inconsiderable latent effect on the incubation period as well as the percentage of hatching of the resulting eggs. A. calamus oil decreased the percentage of egg hatchability with serious increase in the percent sterility. The two tested oils had slight extension in either larval or pupal stage of the resulting progeny of C. maculatus. Both oils show oviposition deterrency towards C. maculatus adults. The oviposition deterrent index was 100% when mung bean seeds were treated with A. calamus oil. The bruchid females laid all eggs on control seeds and no eggs were laid on treated seeds.     

Nest construction and larval behaviour of Bubas bison (L.) and Bubas bubalus (01.) (Coleoptera, Scarabaeidae)

Abstract.

• 1 Female beetles working alone or in cooperation with a male excavated vertical, tunnel-shaped brood chambers. Each chamber was filled with dung to form a cylindrical brood mass which contained two eggs, one near each pole.
• 2 To examine the possible relationship with other Onitini (which lay either one or several eggs per brood mass) factors that influence the two-egg programme were studied. Brood masses with only a single egg were formed if excavation was resumed prematurely. Conversely, when excavation was suppressed several oviposition programmes fused to produce a multi-egg brood mass.
• 3 The larvae repaired their chambers in the typical Scarabaeine manner by building a self-supporting wall formed from their own excrement. This behaviour also prevented direct contact and fighting between adjacent larvae in the same brood mass, and it allowed the larvae to survive inside artificial brood balls. Similar behaviour was observed in larvae of Onthophagus taunts and Ontho-phagus vacca (which develop in one-egg brood masses). The evolution of nesting habits that involve multi-egg brood masses or free-standing brood balls may depend on the pre-existence of this larval repair behaviour.

     

Oviposition in three species of limnephiloid caddisflies (Trichoptera): hierarchical influences on site selection

1. Oviposition site selection was studied in three trichopteran species; an undescribed species of Hydatophylax (Limnephilidae), Onocosmoecus unicolor (Limnephilidae) and Neophylax rickeri (Uenoidae), in two Coastal Range streams in California, U.S.A. 2. Hydatophylax sp. egg masses were generally found at or above the water surface on substrata near the bank in pools, where undercut banks, overhanging vegetation and rocks shaded attachment sites and provided wind protection. Onocosmoecus unicolor females deposited their egg masses exclusively above the water on moist emergent wood. The egg masses of N. rickeri were almost exclusively found in high velocity areas of riffles, where females oviposited under water and attached their egg masses to the submersed undersides of unembedded, protruding stones with large emergent surfaces. 3. For Hydatophylax sp. and N. rickeri, a hierarchical selection scheme is proposed in which females use cues at three different spatial levels (stream, stream subunit, substratum) to choose oviposition sites. Females of O. unicolor seemed to choose at only two different levels of spatial resolution (stream, substratum). 4. The formation of large aggregations of egg masses in N. rickeri and O. unicolor suggests that females actively choose oviposition sites where conspecific egg masses are already attached. The clustering of egg masses may minimise the risk of predation through the dilution effect, because egg masses of N. rickeri and O. unicolor are both attacked by dipteran predators. 5. Females of the three Trichoptera species studied are able to make a clear choice of oviposition site in the heterogeneous stream environment. In comparison with other stream microhabitats, these are characterised by stable and relatively predictable environmental conditions during the time of egg development, which is seen as the main selective pressure leading to the observed egg‐laying behaviour.     

Oviposition behaviour of the soil mite Pergamasus brevicornis (Acari: Parasitidae)

We observed the oviposition behaviour of the soil mite Pergamasus brevicornis Berlese (Acari: Parasitidae) using continuous video-monitoring. Oviposition consisted of six sequential phases. The first phase (I) involved inspection of the substrate. In the second phase (II) there were rhythmic movements of the first pair of legs and slight reciprocating movements of the body. The third (III) was a resting phase. In the fourth phase (IV) the gnathosoma was lowered and the body was raised. In the next phase (V) there were two sub-phases. During the first (Va), the female held the egg below the gnathosoma. In the second sub-phase (Vb), the gnathosoma moved up holding the egg, which was then placed on the substrate. The last phase (VI) involved intense ‘cleaning’ movements of the chelicerae and palps. During Va a protective external eggshell structure is gradually formed, involving a phase where the egg shell is sticky. After moving the egg to the substrate, the female freed her palps and chelicerae from the sticky egg shell and cleaned her gnathosomal appendages. Phases II–V took on average 207 ± 69 s.     

Environmental adaptation and genetic variations in geographically isolated Emma field crickets Teleogryllus emma (Orthoptera: Gryllidae)

The Emma field cricket, Teleogryllus emma (Ohmachi & Matsuura), distributed between 43°N and 30°N in the Japanese archipelago, is univoltine and overwinters in the egg stage. Its eggs hatch on the slope of the Oishi Dam (38.03°N, 139.57°E, 160–170 m a.s.l.) in late June, adults begin emerging from late August, and oviposition lasts until early October. Oviposition is limited to the period when the water level of the Oishi Dam is low. The period from egg hatching to adult emergence is approximately 1 month shorter than that of the T. emma population on the Arakawa riverside (38.09°N, 139.57°E, 29 m a.s.l.), which is approximately only 7 km from the Oishi Dam. The egg and body sizes of T. emma on the slope of the Oishi Dam were smaller than those of T. emma on the Arakawa riverside, and the egg and nymphal periods were shorter; these variations were inherited by the next generation of T. emma. The egg period, nymphal period and head width of T. emma on the dam slope correspond to those of the populations near 40°N. Several traits of the T. emma population on the dam slope were naturally selected by adapting to the isolated environment, resulting in the genetic variations. However, their variations were small and the period after isolation is short, suggesting that it is in the early stages of speciation.     

Density-dependent egg dispersion in flowers of Silene vulgaris by the seed predator Hadena confusa (Noctuidae)

Abstract.

• 1 Hadena moths drink nectar from, pollinate, and oviposit into the flowers of the genus Silene. In the gynodioecious Silene vulgaris (Moench) Garcke (Caryophyllaceae), eggs were laid by Hadena confusa (Hufn.) females both in male phase flowers, which could not be pollinated by the moth at the time of oviposition, and in female phase and female flowers that allowed pollination.
• 2 Flowers in which an egg had been laid had a lowered probability of receiving a second egg laid by another female during the same night.
• 3 The degree of flower discrimination by the moth with respect to the presence of eggs was positively correlated with the proportion of flowers containing eggs.
• 4 The oviposition behaviour of H.confusa is moulded by four factors: (1) a high probability of chosen male phase flowers later becoming pollinated by other moths, (2) the presence of conspecific eggs and risk for larval competition, (3) probability of presence of conspecific eggs also in neighbouring flowers, (4) capability of larvae to move to adjacent flowers.

     

Repellency and toxicity of extract from Francoeria crispa (Forsk.) to Eutetranychus orientalis (Klein) (Acari: Tetranychidae)

Laboratory bioassays were conducted to characterise the activity of the extract Francoeria crispa (Forsk.) (Family: Compositae) against the citrus brown mite, Eutetranychus orientalis (Klein). Ethyl acetate was tested for preparing the crude extract of F. crispa. The extract was tested for its toxicity against eggs and adult females of the pest E. orientalis. Ethyl acetate extract of F. crispa affected the behaviour, toxicity and fecundity of females under laboratory conditions. The extract had similar toxic effects on egg stage and adult females of E. orientalis (LC₅₀ = 0.00050 g/ml), respectively. Leaf discs treated with increasing concentrations of ethyl acetate extract of F. crispa showed a high percentage of repellency (97.45%), respectively. Treated females with LC₅₀ concentration of ethyl acetate extract showed a higher remarkable percentage of mortality as well as a reduction in the total number of eggs laid during 7 days. Ten isolated fractions of ethyl acetate crude extract from F. crispa were detected. The results clearly indicate that the isolate number (10) was the most toxic isolate on eggs and females of E. orientalis (LC₅₀ = 0.00014 and 0.000125 g/ml), respectively.     

Characterisation of the arrestment responses of Trichogramma evanescens

Summary Contact kairomones and oviposition in a host egg stimulated arrestment behaviour in Trichogramma evanescens, characterised by a reduction in walking speed and increased turning. Previous oviposition experience did not influence a parasitoid's response to contact kairomones, but successive encounters with kairomone patches resulted in parasitoids habituating to the contact chemical. Oviposition on a kairomone patch did not reverse this habituation effect. It was concluded that contact kairomones and host eggs will both contribute independently to the duration of a patch visit. The selection of patches by T. evanescens will depend on its response to kairomones. Results from this study indicate that the application of contact kairomones to field crops will not necessarily increase the probability of parasitoids finding hosts.     

Seasonal synchrony of life cycles between the exotic predator,Pseudoscymnus tsugae (Coleoptera: Coccinellidae) and its prey,the hemlock woolly adelgid Adelges tsugae (Homoptera:Adelgidae)

1 The seasonal synchrony between the exotic predator, Pseudoscymnus tsugae and its prey, the hemlock woolly adelgid, Adelges tsugae, was investigated in field cages and in the forest in Connecticut, U.S.A. from 1997–1999. 2 In early spring, egg to adult development took 45 d at 18.7 °C, 39.7 d at 20.2 °C and 31.5 d at 22.7 °C. Earliest emerging F1 adults mated and oviposited in the same year. whereas F1 and F2 females emerging later in the summer mated and reserved most of their egg complement for the following year. 3 A second generation of P. tsugae is possible in Connecticut but may be delayed by cool mid‐spring temperatures. Individuals of three generations of P. tsugae, including overwintering survivors, may coexist in July and August and adults can be found year‐round with A. tsugae in infested hemlock forests. 4 A linear regression model for development from egg to adult under field temperatures gave good agreement with results from constant temperature findings. The model predicted a lower development threshold of 9.5 °C and a sum of effective temperatures of 405 day °C. Development time of P. tsugae is shorter relative to its prey A. tsugae and generation time ratios of predator to prey was 0.16–0.5, with an advantage conferred on the coccinellid. 5 Overwintering ability and behaviour were determined in 1998–1999 and adults remained on infested hemlock branches throughout a mild winter, becoming reproductively active in mid‐April. Peak oviposition period extended from April to July, in synchrony with peak oviposition and developing stages of two generations of A. tsugae.     

Complete development on Elodea canadensis (Hydrocharitaceae) eliminates Hydrellia egeriae (Diptera,Ephydridae) as a candidate biological control agent of Egeria densa (Hydrocharitaceae) in the U.S.A.

The leaf-mining fly Hydrellia egeriae Rodrigues-Junior (Diptera: Ephydridae) was evaluated under quarantine conditions as a candidate biological control agent of the invasive aquatic weed Egeria densa Planchon (Hydrocharitaceae) in California, U.S.A. The objective of this study was to test the fly’s feeding preference for E. densa as compared to the North American native Elodea canadensis Michaux (Hydrocharitaceae), an important benchmark species. No choice testing revealed that H. egeriae can complete development on E. canadensis and choice testing indicated the fly presents little oviposition discrimination between E. densa and E. canadensis. It is concluded that the host range of H. egeriae is too broad for use as a biological control agent of E. densa in the U.S.A.     

Behavioural plasticity in relation to egg and time limitation: the case of two fly species in the genus Anastrepha (Diptera: Tephritidae)

Reproductive opportunities in insects that deposit their eggs in discrete resource patches are frequently limited because the availability of oviposition substrates is often spatially and temporally restricted. Such environmental variability leads individuals to confront time‐ or egg‐limitation constraints. Additionally, species with different oviposition strategies (i.e. single egg layers vs clutch layers) commonly deal with different structural and ecological characteristics of larval host plants. To test the hypothesis that oviposition strategies such as laying eggs singly or in batches (clutches) are related to these constraints (i.e. egg vs time limitation), we compared the lifetime oviposition patterns of two closely related sympatric species of Anastrepha (Diptera: Tephritidae) with different oviposition strategies. We exposed five cohorts of A. obliqua and A. ludens females, over the course of their adult lifetimes, to three conditions of “habitat quality” (measured as host density per cage): unpredictable habitat quality (host density varied randomly from day to day between 1, 5, 15, 30 and 60 hosts/cage), low habitat quality (fixed density of one host/cage) and high habitat quality (fixed density of 60 hosts/cage).
Responses to host density conditions were strikingly different in the two species. (1) Frequency of host visits and oviposition events increased in A. obliqua but not in A. ludens when host densities increased. (2) Anastrepha ludens females accepted low quality hosts (i.e. fruits on which eggs had already been laid and were therefore partially covered with host marking pheromone) significantly more often than A. obliqua females did. (3) Females of A. obliqua adjusted their oviposition activity to variations in host density, whereas A. ludens females exhibited a constant oviposition pattern (i.e. did not respond to variations in host density). Based on the above, it is likely that in A. obliqua oviposition is governed by egg‐limitation and in A. ludens by time‐limitation constraints. We discuss the relationship between the oviposition strategies of each fly species and the fruiting phenology and density of their native host plants. We also address the possible influence of oogenesis modality and parasitism by braconid wasps in shaping oviposition behaviour in these insects.