Host-seeking behavior of Ixodes pacificus (Acari: Ixodidae) nymphs in relation to environmental parameters in dense-woodland and woodland-grass habitats

In the far-western United States, the bacteria that cause Lyme disease (Borrelia burgdorferi, Bb) and human granulocytic anaplasmosis (Anaplasma phagocytophilum, Ap) are transmitted by the western black-legged tick (Ixodes pacificus). In a dense woodland, human behaviors involving contact with wood were recently found to pose greater risk for encountering I. pacificus nymphs than behaviors entailing exclusive exposure to leaf litter. A four-year follow-up study was undertaken in the same woodland and, as a comparison area in one year only, in a nearby woodland-grass habitat to explore the biotic and abiotic factors that might elevate human exposure to host-seeking nymphs. Nymphs were active in the dense woodland throughout the daytime, but no consistent pattern of activity was observed with respect to time of day, temperature, or relative humidity. Significantly more nymphs were collected from the southern aspects of dense-woodland trunks than from other aspects, and more nymphs quested at a height of < or = 1 m vs 1-2 m aboveground. The prevalence of bacterial infection in ticks from the dense woodland was highly variable among years, with maxima of 22.6% and 42.9% for Bb, and 15.6% and 1.8% for Ap, in nymphs from logs and trunks, respectively. The mean densities of nymphs, and of Bb- or Ap-infected nymphs, were typically higher on logs and trunks than in adjoining leaf litter or grass in both habitats. The acarologic risk of encountering an infected nymph on dense-woodland logs or trunks was 2.8 to 11 times higher for Bb than for Ap in two of three years, and it was usually higher in dense woodland than in woodland-grass for both agents. Coinfections were rare (0.27%, n = 369 nymphs from both habitats). Individuals having prolonged contact with logs or trunks in spring would be well advised to employ personal protective measures to minimize exposure to I. pacificus nymphs and their attendant bacterial zoonotic agents.     

Dispersal Capacity and Behavior of Nymphal Stages of Halyomorpha halys (Hemiptera: Pentatomidae) Evaluated Under Laboratory and Field Conditions

The invasive brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), is a highly polyphagous and mobile pest causing crop damage aggregated at the perimeters of crop fields. Understanding the dispersal biology of H. halys is critical for the development of reliable monitoring and management strategies. In this study, dispersal ecology of H. halys nymphs was studied under laboratory and field conditions. In the laboratory, horizontal and vertical walking capacity was quantified for mobile nymphal stages (i.e., 2nd through 5th instars) and compared with adults. There was a significant difference in the horizontal distance moved by H. halys among the life stages tested. Third instars exhibited significantly greater walking distances compared with adults; horizontal walking distances by other nymphal stages were not significantly different from adults. A similar pattern was observed from vertical climbing tests of H. halys. Third and 4th instars climbed significantly greater distances compared with 2nd instars and adults, while distances climbed by 5th instars were intermediate. In the field, the walking distance of 3rd and 5th instar nymphs on mowed grass was quantified based on direct observation of individuals for 30 min. Under these conditions, 5th instars moved nearly two-fold greater distances compared with 3rd instars, but surface temperature affected both nymphal stages similarly. Shorter bouts of movement were common at surface temperatures below 25 °C, whereas individuals showed longer walking distances above 25 °C. In mark-release-recapture studies, 4th and 5th instars were released and recaptured in traps baited with attractive pheromonal-based stimuli to estimate dispersal rates under field conditions. When insects were released 5 m from traps, both instars were first recaptured within 2 h after release, with the recapture rates of 54 and 69 % for 4th and 5th instars over 24 h, respectively. When insects were released 20 m from traps, 4th and 5th instars were first recaptured in less than 5 h, with the recapture rates of 27 and 51 %, respectively. The results of this study indicate that H. halys nymphs have strong dispersal capacity with which populations can easily move among host plants and other attractive stimuli at farmscape levels.     

Risk indicators for the tick Ixodes ricinus and Borrelia burgdorferi sensu lato in Sweden

The distributional area of the tick Ixodes ricinus (L.), the primary European vector to humans of Lyme borreliosis spirochaetes (Borrelia burgdorferi sensu lato) and tick‐borne encephalitis virus, appears to be increasing in Sweden. It is therefore important to determine which environmental factors are most useful to assess risk of human exposure to this tick and its associated pathogens. The geographical distribution of I. ricinus in Sweden was analysed with respect to vegetation zones and climate. The northern limit of I. ricinus and B. burgdorferi s.l. in Sweden corresponds roughly to the northern limit of the southern boreal vegetation zone, and is characterized climatically by snow cover for a mean duration of 150 days and a vegetation period averaging 170 days. The zoogeographical distribution of I. ricinus in Sweden can be classified as southerly–central, with the centre of the distribution south of the Limes Norrlandicus. Ixodes ricinus nymphs from 13 localities in different parts of Sweden were examined for the presence of B. burgdorferi s.l. and found to be infected with Borrelia afzelii and Borrelia garinii. Tick sampling localities were characterized on the basis of the density of Borrelia‐infected I. ricinus nymphs, presence of specific mammals, dominant vegetation and climate. Densities of I. ricinus nymphs and Borrelia‐infected nymphs were significantly correlated, and nymphal density can thus serve as a general indicator of risk for exposure to Lyme borreliosis spirochaetes. Analysis of data from this and other studies suggests that high densities of Borrelia‐infected nymphs typically occur in coastal, broadleaf vegetation and in mixed deciduous/spruce vegetation in southern Sweden. Ixodes ricinus populations consistently infected with B. burgdorferi s.l. can occur in: (a) biotopes with shrews, rodents, hares and birds; (b) biotopes with shrews, rodents, hares, deer and birds, and (c) island locations where the varying hare (Lepus timidus) is the only mammalian tick host.     

Relative importance of lizards and mammals as hosts for ixodid ticks in northern California

Lizards and mammals were trapped and examined for ticks from August 1992 to June 1993 in two habitat types, chaparral and woodland-grass, in northern California. Five tick species were collected from mammals (Dermacentor occidentalis, Haemaphysalis leporispalustris, Ixodes pacificus, I. spinipalpis, I. woodi), but only I. pacificus was found on lizards. Dermacentor occidentalis, I. pacificus, and I. woodi occurred in both habitats, whereas H. leporispalustris and I. spinipalpis were found only on animals trapped in chaparral. The tick species most commonly encountered on mammals was D. occidentalis in chaparral and I. pacificus in woodland-grass. Peak infestation of mammals occurred in spring for I. pacificus immatures and H. leporispalustris, summer for D. occidentalis immatures, fall through spring for I. woodi immatures, and fall through winter for I. spinipalpis. The primary aim of the study was to quantify the relative importance of the western fence lizard (Sceloporus occidentalis), which is reservoir-incompetent for Borrelia burgdorferi sensu lato (s.l.), and mammalian B.burgdorferi s.l.-reservoirs as hosts for the immature stages of I. pacificus in spring. The estimated relative utilization by I. pacificus of the western fence lizard versus mammals was 88% for larvae and 99% for nymphs in chaparral in May. When tick infestation data were corrected for a two-fold lower efficiency of field examinations for rodents than for lizards, the western fence lizard still accounted for 78% of larval and 98% of nymphal feedings. In woodland-grass, 46% of 100 I. pacificus larvae and 100% of 52 nymphs recovered from mammals or western fence lizards during May-June were collected from the lizards. However, this may represent an underestimate of the importance of the western fence lizard as a larval host in this habitat because inclement weather during the late May sampling period doubtless resulted in significantly decreased lizard activity. In conclusion, the western fence lizard was more heavily utilized by I. pacificus immatures, especially nymphs, than were rodents. This revised version was published online in July 2006 with corrections to the Cover Date.     

Dispersal of the cabbage root fly

The dispersal rates of wild and culture cabbage root flies Erioischia brassicae were determined in release-recapture experiments at Wellesbourne in 1971–3. The experiments were concerned mainly with the first 7 days of adult life. The flies, released from nine locations in the area, were recaptured in yellow water-traps. Dispersal was affected by wind, rain and the terrain the flies were crossing. The flies least often recaptured were those released into the host crop when 6–12 days old. The results indicated the following pattern of behaviour. Flies moved little during the first 2 days of adult life but by the third day both sexes had dispersed to c. 100 m from the release point. Flies are known to mate about the fourth day and after this the males continued to disperse at c. 100 m per day for the three subsequent days. ‘Wild’ females from field-collected pupae carried out a ‘migratory’ flight, however, and dispersed at c. 1000 m per day during the fifth and sixth days, the days preceding the start of oviposition. Similar rates of dispersal were recorded from flies released across host crop and non-host crop areas. Some females did not stop at the first crop they encountered. The culture females from the laboratory-reared pupae dispersed only c. one-third of the distance of the wild females. There was considerable intermingling of local populations. The percentage recapture of young culture and wild flies released during the pre-oviposition period of this species was 38 ± 4 and 19 ±4 for males, and 15 + 2 and 8+1 for females, respectively. The dispersal range of the cabbage root fly is probably within a 2000–3000 m radius of the site of infestation.     

Development and comparison of trunk traps to monitor movement of Halyomorpha halys nymphs on host trees

Halyomorpha halys Stål (Hemiptera: Pentatomidae) has recently become a major orchard pest in the Mid‐Atlantic, USA. Large H. halys populations can develop on wild tree hosts adjacent to orchards, posing an ongoing threat to fruit. Adults and nymphs feed on tree fruit, causing economic injury. Understanding the seasonal patterns of nymphal host use among trees at the orchard‐woodland interface may aid the development of integrated pest management strategies for this pest. In laboratory and field experiments, modified versions of published trap designs – ‘Circle’, ‘Hanula’, ‘M&M’ (after Moeed & Meads) traps – were compared for their effectiveness for capturing H. halys nymphs walking up and down tree trunks. In the laboratory, second instars were released at the top and bottom of ailanthus (tree of heaven), Ailanthus altissima (Mill.) Swingle (Simaroubaceae), logs and captures were recorded after 24 h. Circle and M&M traps, respectively, were most effective for capturing nymphs walking up and down. In the field, traps were deployed on ailanthus trees next to apple orchards and captures were recorded weekly from 24 July to 11 September 2013. As in the laboratory, Circle and M&M traps captured the greatest number of upward‐ and downward‐walking nymphs. Hanula traps were least effective in both experiments. In the field, 88% of total captures were of nymphs walking up trees. This was at least partially explained by behavioral assays in the laboratory demonstrating that nymphs exhibited negative gravitaxis and positive phototaxis. Stage‐specific trends in captures of instars walking up during field sampling were observed. These results suggest that trunk traps can be used to address important ecological questions about seasonal patterns of host use by H. halys nymphs.     

Influence of harmonic radar tag attachment on nymphal Halyomorpha halys mobility,survivorship, and detectability

The brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), is a polyphagous invasive insect and currently one of the most threatening agricultural pests in the USA and globally. Nymphs are highly mobile, moving among host plants, and causing significant damage. Thus, understanding dispersal biology for all life stages is critical for the development of reliable monitoring and management programs. Here, we evaluated the influence of harmonic radar as a tool to study dispersal ecology of nymphal H. halys; we measured the impact of glues and tag attachment on survivorship and mobility in the laboratory and validated in the field that tagged and released nymphs could be tracked on baited and unbaited host and non‐host plants using harmonic radar. In the laboratory, four glues were evaluated for attaching harmonic radar tags securely to nymphs, and survivorship with attached tags was measured. There were no significant differences in survivorship or vertical and horizontal movement among nymphs with tags affixed with the glue treatments compared with the untagged control. Based on numerically greater survivorship of nymphs with tags affixed with Loctite glass glue, a field validation study of tagged nymphs released in host (apple tree) and non‐host (mowed grass) with or without H. halys pheromonal stimuli present revealed that nymphs could be successfully relocated using harmonic radar after 48 h. Among treatments, 83% of nymphs remained in baited and unbaited apple trees, 50% of nymphs remained in baited mowed grass plots, and in unbaited mowed grass plots, 17% of fifth instars, and 0% of fourth instars were retained. The absence of negative effects on mobility, survivorship, and field tracking validates that harmonic radar can be used to study dispersal ecology of nymphal H. halys.     

Survival rates of immature Ixodes pacificus (Acari: Ixodidae) ticks estimated using field‐placed enclosures

Granulocytic anaplasmosis (GA) and Lyme borreliosis are emerging tick‐borne diseases caused by infection with Anaplasma phagocytophilum and Borrelia burgdorferi, respectively, and maintained in rodent‐Ixodes spp. tick cycles, including I. pacificus in the western U.S. Ixodes pacificus has a multiple‐year life cycle and B. burgdorferi and A. phagocytophilum are transstadially, but not transovarially, transmitted within ticks, thus ticks function importantly in maintaining infection in nature. In this study, the survival of larval and nymphal I. pacificus was determined using ticks placed in tubes in leaf litter from June 2005 to September 2006 at two field sites in the California northern coast range mountains and a laboratory control. In all three sites, nymphal and larval survival ranged from 90–400 d, with differences in mean survival among sites. Fewer ticks died in the autumn in the moister field sites compared with the drier incubator control treatment. The first large die‐off event in late autumn occurred at all sites shortly before relative humidity increased from 80–100% and temperature declined from approximately 22–15° C. The concurrent die‐off in the incubator population, subject to relative humidity and temperature regimes that were invariant, suggests that survival time was dependent on other factors in addition to environmental conditions. These results suggested that many ticks exhausted resources or tolerance for relatively low humidity within six months of questing, and that higher humidity prolonged survival. Based on observed longevity, humans and other animals could acquire A. phagocytophilum infection from adult I. pacificus that were infected up to three years earlier.     

Host odour composition affects host location efficiency of tsetse (Diptera, Glossinidae)

Abstract. Marked Glossina pallidipes Austen were released downwind of an odour source in the field in Zimbabwe and the percentage recaptured at the source on the same day was measured.In the absence of odour, 1.3% of the marked tsetse released from a box or refuge were recaptured, independent of the distance between release point and odour source.The distance was varied from 10 to 100 m.When natural ox odour or a blend of carbon dioxide, acetone, octenol and phenols was dispensed, untransformed recapture percentages of box-released tsetse decreased from 18% for tsetse released at 10 m to 2% for tsetse released at 100 m.Recapture percentages were significantly higher than in the absence of odour at all release distances for ox odour and for release distances up to 75 m downwind for the artificial odour.When a combination of acetone, octenol and phenols or carbon dioxide on its own was dispensed, recapture percentages decreased from 6% for tsetse released at 10 m to 0% for tsetse released at 100 m.With these odours, recapture percentages were higher than in the absence of odour when tsetse were released at 20 m from the source, but were lower than recaptures in the presence of ox odour or the artificial mixture with carbon dioxide.Recapture percentages of flies spontaneously leaving refuges were higher than those of box-released tsetse.Proximity of source had no effect on the recapture percentage of refuge-leaving tsetse and host-location efficiency was close to 100% when host odour was detected at 30 m or less.The results are discussed in relation to the host location strategy of tsetse.     

Field and climate‐based model for predicting the density of host‐seeking nymphal Ixodes scapularis,an important vector of tick‐borne disease agents in the eastern United States

Aim Ixodes scapularis is the most important vector of human tick‐borne pathogens in the United States, which include the agents of Lyme disease, human babesiosis and human anaplasmosis, among others. The density of host‐seeking I. scapularis nymphs is an important component of human risk for acquiring Borrelia burgdorferi, the aetiological agent of Lyme disease. In this study we used climate and field sampling data to generate a predictive map of the density of host‐seeking I. scapularis nymphs that can be used by the public, physicians and public health agencies to assist with the diagnosis and reporting of disease, and to better target disease prevention and control efforts. Location Eastern United States of America. Methods We sampled host‐seeking I. scapularis nymphs in 304 locations uniformly distributed east of the 100th meridian between 2004 and 2006. Between May and September, 1000 m² were drag sampled three to six times per site. We developed a zero‐inflated negative binomial model to predict the density of host‐seeking I. scapularis nymphs based on altitude, interpolated weather station and remotely sensed data. Results Variables that had the strongest relationship with nymphal density were altitude, monthly mean vapour pressure deficit and spatial autocorrelation. Forest fragmentation and soil texture were not predictive. The best‐fit model identified two main foci – the north‐east and upper Midwest – and predicted the presence and absence of I. scapularis nymphs with 82% accuracy, with 89% sensitivity and 82% specificity. Areas of concordance and discordance with previous studies were discussed. Areas with high predicted but low observed densities of host‐seeking nymphs were identified as potential expansion fronts. Main conclusions This model is unique in its extensive and unbiased field sampling effort, allowing for an accurate delineation of the density of host‐seeking I. scapularis nymphs, an important component of human risk of infection for B. burgdorferi and other I. scapularis‐borne pathogens.     

Notes on paternal care and sibling cannibalism in the giant water bug, Lethocerus deyrolli (Heteroptera: Belostomatidae)

Males of the giant water bug, Lethocerus deyrolli, care for egg masses on vegetation above the water surface. They supply the developing eggs with water and guard them against predators. In the present study, mechanisms by which paternal care is extended were found. Males were found situated just below the water on the natal substrate (usually a stick), and the first instar nymphs were aggregated around the substrate. When disturbed, the males showed aggressive behavior, threatening the intruder with their forelegs. Nymphs up to 12 h old did not attack the offered sibling nymphs or anuran larvae, which are common prey in the field. The 24 h‐old nymphs attacked both prey animals; however, they preferred anuran larvae. Cannibalistic behavior in the nymphs was well developed 72 h after hatching, when the nymphs had already dispersed from the natal substrate. The suppression of sibling cannibalism in younger nymphs would promote the maintenance of tight nymphal aggregations and consequently extend male care in this predatory species.     

Detection of Lyme disease spirochete,Borrelia burgdorferi sensu lato,including three novel genotypes in ticks (Acari: Ixodidae) collected from songbirds (Passeriformes) across Canada

Lyme disease is reported across Canada, but pinpointing the source of infection has been problematic. In this three‐year, bird‐tick‐pathogen study (2004–2006), 366 ticks representing 12 species were collected from 151 songbirds (31 passerine species/subspecies) at 16 locations Canada‐wide. Of the 167 ticks/pools tested, 19 (11.4%) were infected with Borrelia burgdorferi sensu lato (s.l.). Sequencing of the rrf‐rrl intergenic spacer gene revealed four Borrelia genotypes: B. burgdorferi sensu stricto (s.s.) and three novel genotypes (BC genotype 1, BC genotype 2, BC genotype 3). All four genotypes were detected in spirochete‐infected Ixodes auritulus (females, nymphs, larvae) suggesting this tick species is a vector for B. burgdorferi s.l. We provide first‐time records for: ticks in the Yukon (north of 60° latitude), northernmost collection of Amblyomma americanum in North America, and Amblyomma imitator in Canada. First reports of bird‐derived ticks infected with B. burgdorferi s.l. include: live culture of spirochetes from Ixodes pacificus (nymph) plus detection in I. auritulus nymphs, Ixodes scapularis in New Brunswick, and an I. scapularis larva in Canada. We provide the first account of B. burgdorferi s. l. in an Ixodes muris tick collected from a songbird anywhere. Congruent with previous data for the American Robin, we suggest that the Common Yellowthroat, Golden‐crowned Sparrow, Song Sparrow, and Swainson's Thrush are reservoir‐competent hosts. Song Sparrows, the predominant hosts, were parasitized by I. auritulus harboring all four Borrelia genotypes. Our results show that songbirds import B. burgdorferi s.l.‐infected ticks into Canada. Bird‐feeding I. scapularis subadults were infected with Lyme spirochetes during both spring and fall migration in eastern Canada. Because songbirds disperse millions of infected ticks across Canada, people and domestic animals contract Lyme disease outside of the known and expected range.     

Return to preferred habitats (edges) as a function of distance in Akodon azarae (Rodentia, Muridae) in cropfield-edge systems of central Argentina

We studied the ability of Akodon azarae (Rodentia, Muridae) to return to their preferred habitat, when released at a perpendicular distance (25, 50 or 75 m) from the edge towards the cropfield, and a parallel distance (100 m) away from the site of first capture within the edge habitat. Return success was estimated as the proportion of animals recovered in edges. The recapture rate between the field and the border was significantly higher than the recapture rate estimated according to successive captures in the border. Successful returns did not decrease significantly with increasing release distance, but animals released at 50 m from edges were less successful in returning to borders than the other release-distance groups. Although the median time taken to the first recapture in edges did not differ among the release-distance groups, rodents released at 25 m and 50 m returned to edges faster than those released at 75 m. A. azarae showed both a successful return to the edge and a trend to return to the home range area. We conclude that A. azarae can return to edges from cropfields at distances that are larger than those they usually travel, allowing the use of fields when they present good conditions for reproduction and survival. Successful return is probably the result of direct movements rather than random wandering.     

Transmission cycles of Borrelia burgdorferi and B. bissettii in relation to habitat type in northwestern California

This study was undertaken to determine which rodent species serve as primary reservoirs for the Lyme disease spirochete Borrelia burgdorferi in commonly occurring woodland types in inland areas of northwestern California, and to examine whether chaparral or grassland serve as source habitats for dispersal of B. burgdorferi‐ or B. bissettii‐infected rodents into adjacent woodlands. The western gray squirrel (Sciurus griseus) was commonly infected with B. burgdorferi in oak woodlands, whereas examination of 30 dusky‐footed woodrats (Neotoma fuscipes) and 280 Peromyscus spp. mice from 13 widely‐spaced Mendocino County woodlands during 2002 and 2003 yielded only one infected woodrat and one infected deer mouse (P. maniculatus). These data suggest that western gray squirrels account for the majority of production by rodents of fed Ixodes pacificus larvae infected with B. burgdorferi in the woodlands sampled. Infections with B. burgdorferi also were rare in woodrats (0/47, 0/3) and mice (3/66, 1/6) captured in chaparral and grassland, respectively, and therefore these habitats are unlikely sources for dispersal of this spirochete into adjacent woodlands. On the other hand, B. bissettii was commonly detected in both woodrats (22/47) and mice (15/66) in chaparral. We conclude that the data from this and previous studies in northwestern California are suggestive of a pattern where inland oak‐woodland habitats harbor a B. burgdorferi transmission cycle driven primarily by I. pacificus and western gray squirrels, whereas chaparral habitats contain a B. bissettii transmission cycle perpetuated largely by I. spinipalpis, woodrats, and Peromyscus mice. The dominant role of western gray squirrels as reservoirs of B. burgdorferi in certain woodlands offers intriguing opportunities for preventing Lyme disease by targeting these animals by means of either host‐targeted acaricides or oral vaccination against B. burgdorferi.     

Relationship between temporal abundance of ticks and incidence of Lyme borreliosis in Lower Silesia regions of Poland

The aim of this study was to identify the factors determining the incidence of Lyme borreliosis (LB) in south‐western Poland by estimating the prevalence of B. burgdorferi s. l. in I. ricinus, and to analyze the temporal abundance of ticks in relation to epidemiological data on LB incidence. Host‐seeking ticks collected in 2011 in four districts in southwestern Poland were examined by nested PCR for the presence of B. burgdorferi s.l. In total, 2,507 host‐seeking I. ricinus were collected. The temporal abundance of ticks varied between districts. The minimal infection rates with B. burgdorferi s.l. were 11.5% for nymphs and 37.7% for adults. There were no statistical differences in the level of infection between districts either for nymphs or for adults. Five different genospecies were identified within the B. burgdorferi s.l. complex: B. garinii, B. afzelii, B. lusitaniae, B. valasiana, and B. burgdorferi s.s., and additionally B. miyamotoi. Our results point to a relationship between tick temporal abundance and LB incidence both for adults and nymphs. The high abundance of ticks is positively correlated with the number of LB cases in humans. The tick's abundance may be considered as a major factor in determining the LB risk in southwestern Poland.     

The probing behaviour of nymphs of Vanduzeea arquata and Enchenopa binotata (Homoptera: Membracidae) on host and non-host plants

Abstract. 1. Nymphs of Vanduzeea arquata Say have been found to be more host-specific in nature and to show a higher degree of selectivity in host discrimination experiments than nymphs of Enchenopa binotata (Say), It was hypothesized that this differential selectivity would be reflected in the probing behaviour of individuals placed on twigs of host and non-host plants. Probing behaviour was examined by direct observation of nymphs and by sectioning and staining the probed plant tissues.
2. All nymphs probed readily and for extended periods on both host and non-host twigs. E.binotuta nymphs showed no consistent differences in probing behaviour on hosts versus non-hosts, but V.atquuta nymphs were more likely to withdraw their stylets within 60 s when on non-host twigs and produced honeydew only when on their host species. V.urquatu nymphs reached the phloem sieve elements only when on host twigs and broke many cells in peripheral plant tissue layers while probing. E.binotata nymphs broke few cells and often reached the phloem of non-host as well as host plants.
3. Nymphs of V.arquata always reject non-host plants, apparently in the course of probing and prior to encountering the phloem sap. Chemical compounds released from ruptured parenchyma cells may act as probing stimulants or inhibitors. E.binotura nymphs often feed on non-host plants in a non-choice situation; their preferential settling on host twigs in discrimination experiments may reflect a tendency to abandon non-host twigs more readily than host twigs.     

Measurements of locomotor activity in hatchlings of the migratory locust Locusta migratoria: effects of intrinsic and extrinsic factors

Abstract. The effects of intrinsic and extrinsic factors on locomotor activity in crowd‐reared first‐stadium nymphs of the migratory locust Locusta migratoria are investigated under continuous light conditions using an actograph apparatus. Nymphs show monomodal or bimodal patterns of locomotor activity with respect to the time after the start of measurements, depending on the age. Locomotor activity is suppressed by the presence of grass in nymphs aged 0–2 days old but a peak of activity observed shortly after hatching is not suppressed. The results suggest that newly‐hatched nymphs may try to disperse from the hatching site. Nymphs show higher locomotor activity levels under moist conditions than under dry conditions during the first 5‐h period of measurements. This enhanced locomotor activity may constitute attempts to avoid high humidity. Under dim‐light conditions (2 × 10^?2 Wm^?2), locomotor activity is suppressed during the first half day and increases to a high level thereafter in both grass‐fed and unfed individuals. This increased activity might indicate a possible involvement of circadian rhythms. Background colour has no significant effect on the locomotor activity. The present study provides new aspects of behaviour in nymphs as well as baseline data for behavioural analysis of locust locomotion in relation to phase polyphenism.     

A comparison of the attraction of nymphs and adults of the ticksAmblyomma hebraeum andA. variegatum to carbon dioxide and the male-produced aggregation-attachment pheromone

The attraction of nymphs of the ticksAmblyomma hebraeum andA. variegatum to CO₂ alone and CO₂ together with the male-produced aggregation-attachment pheromone (AAP) was investigated. Matching experiments on the attraction of the adults of the two species were run for comparison. Nymphs ofA. hebraeum were strongly attracted to sources of CO₂ and CO₂/AAP at distances of 10 and 15 m. At distances of 20 and 25 m significantly more nymphs were attracted to CO₂/AAP than to CO₂ alone. Adults ofA. hebraeum were not attracted to CO₂ alone at any distance; these ticks were attracted to CO₂ and AAP together for distances of up to 25 m. The patterns of attraction of nymphs and adults ofA. variegatum at 10 m were similar to those recorded inA. hebraeum.     

Dispersal of single‐ and double‐brood populations of the European earwig,Forficula auricularia: a mark‐recapture experiment

Quantitative information on dispersal of insects should be taken into consideration for making efficient pest management decisions. Such information was not available for the European earwig, Forficula auricularia L. (Dermaptera: Forficulidae), an important biocontrol agent in fruit orchards. A mark‐recapture experiment was carried out in Belgian orchards, where marked earwigs were released at a single point and recaptured after 1 month. Dispersal from this release point was analysed using an analytical formula of a simple diffusion model with disappearance (e.g., as a result of death) derived by Turchin & Thoeny (1993; Quantifying dispersal of southern pine beetles with mark‐recapture experiments and a diffusion model. Ecological Applications 3: 187) . The cumulative number of recaptured earwigs as a function of the distance of release was used to fit the model and estimate parameters. A derived expression, in terms of these parameters, was used to estimate the frequency distribution of the population, as the radius of a circle enclosing various proportions of the earwigs’ dispersal distances. In Belgium, populations of the European earwig can have two life‐history strategies, single‐ (SBP) and double‐brood populations (DBP). Therefore, mark‐recapture experiments were carried out on both population types. We fitted data from SBP (n = 10) and DBP (n = 16) successfully in both the diffusion model and in an exponential curve. Because of the biological relevance, estimates of the diffusion model were used for calculating the frequency distributions. Males and females dispersed the same distances. No differences were found between orchards with different spatial structures (apple and pear). According to literature data, mobility of earwigs is very low compared with other arthropods, which has consequences for the efficiency of biocontrol interventions, like mass releases of earwigs or the use of hedgerows for the establishment of healthy (source) populations. Quantitative results revealed that earwigs of SBP dispersed four times further than earwigs of double‐brood populations. For instance, 95% of the population remained within a radius of 28.6 m in SBP and 7.54 m in DBP.     

Effect of host deprivation on the foraging behavior of the Asian citrus psyllid parasitoid Tamarixia radiata: observations from the laboratory and the field

The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae), is a significant citrus pest and the parasitoid Tamarixia radiata (Waterston) (Hymenoptera: Eulophidae) has been released in various citrus‐producing areas in classical biological control programs targeting D. citri. We investigated the effect of host deprivation on the foraging behavior and patch utilization by T. radiata. In the laboratory, females deprived of hosts for 3 days tended to leave patches of 12 nymphs without parasitizing hosts during the ca. 30 min they spent in the patch before leaving. Moreover, half of these females failed to host feed, and those that did host feed, on average, needed more than 15 min to complete feeding. Conversely, non‐host‐deprived females parasitized on average three nymphs before leaving patches without host feeding during the ca. 39 min they spent in the patch. These laboratory observations were compared to mass‐reared female T. radiata that were released onto colonies of D. citri nymphs infesting citrus in the field. Release vials were provisioned with honey and these females had no opportunities to host feed over the 1‐ to 2‐day containment period prior to release. When introduced onto D. citri colonies, 68% T. radiata females abandoned D. citri patches prior to probing hosts, in part, because Argentine ants, Linepithema humile (Mayr) (Hymenoptera: Formicidae), tending colonies disturbed searching parasitoids. These results from laboratory and field studies are discussed in the context of classical biological control, with the aim of understanding how to manipulate host availability and ant activity so establishment rates and impact of T. radiata can be improved.