Male-produced aggregation pheromone of Carpophilus sayi, a nitidulid vector of oak wilt disease, and pheromonal comparison with Carpophilus lugubris

Abstract 1 Carpophilus sayi, a nitidulid beetle vector of the oak wilt fungus, Ceratocystis fagacearum, was shown to have a male‐produced aggregation pheromone. 2 Six male‐specific chemicals were identified from collections of volatiles. The two major compounds were (2E,4E,6E,8E)‐3,5‐dimethyl‐7‐ethyl‐2,4,6,8‐undecatetraene and (2E,4E,6E,8E)‐3,5,7‐trimethyl‐2,4,6,8‐undecatetraene, in a ratio of 100 : 18. These compounds, in a similar ratio, were previously reported to be the pheromone of Carpophilus lugubris, a closely related species. The four minor C. sayi compounds (less than 4% as abundant as the first) were also alkyl‐branched hydrocarbons and consisted of two additional tetraenes and two trienes. 3 The pheromone of C. lugubris was re‐examined to refine the comparison with C. sayi, and C. lugubris was found to have the same additional, minor tetraenes as C. sayi, but not the trienes. 4 A synthetic mixture of the two major compounds was behaviourally active for both sexes of C. sayi in oak woodlands in Minnesota. The pheromone was tested in combination with fermenting whole wheat bread dough (a potent synergist of nitidulid pheromones). The combination of the 500‐µg pheromone dose and dough attracted at least 30‐fold more C. sayi than either pheromone or dough by itself. The synergized pheromone has potential as a tool for monitoring insect vector activity in an integrated management program for oak wilt. 5 Although C. lugubris was not present at the Minnesota test sites, two other Carpophilus species, Carpophilus brachypterus and Carpophilus corticinus, were clearly cross‐attracted to the synergized pheromone of C. sayi.     

脊胸露尾甲生物学特性的观察

通过室内饲养观察及实地调查,明确了脊胸露尾甲年发生规律等基本生物学特性。该虫世代重叠现象明显,每代历期约33～38d,其中产卵前期平均68d,卵历期平均32d,幼虫期平均179d,蛹期平均75d。成虫产卵明显受性比的影响,雌雄性比小时,雌虫产卵量增多。     

An attract-and-kill system to control Carpophilus spp. in Australian stone fruit orchards

During two growing seasons, the use of an attract-and-kill system for control of Carpophilus spp. (Coleoptera: Nitidulidae) and the effective range or drawing power of the attract-and-kill stations were examined in stone fruit orchards in the Goulburn Valley, northern Victoria, Australia. Three attract-and-kill stations, baited with synthetic aggregation pheromone plus co-attractant, were placed about 50 m apart in the upwind corner of each treated block 5–6 weeks before the fruit began to ripen. Large numbers of Carpophilus spp. were caught in the attract-and-kill stations immediately after placement. By the time fruit had ripened, the number of Carpophilus spp. caught had decreased greatly. Fruit damage caused by Carpophilus spp. in treated blocks, especially in 2000–2001 season, was almost zero (0.1% and 0.6%) in trees and on the ground, respectively, whereas the damage levels in control blocks were 5.2% and 19.9% in trees and on the ground, respectively. This study indicates that excellent protection of ripening stone fruit may be achieved by using attract-and kill-stations.     

Effect of physiological state and fungal inoculation on chemically modulated host-plant finding by Carpophilus hemipterus and Carpophilus lugubris

Host orientation by Carpophilus hemipterus L. and Carpophilus lugubris Murray (Coleoptera: Nitidulidae) was investigated in a horizontal wind tunnel to ascertain if these species differ in their response to aseptic and fungal-inoculated hosts, and also to determine how age, nutritional status, diel period, and locomotory opportunity affect these behavioral responses. Both species responded to food odors beginning on Day 3 of adult life by walking upwind to the source; flight activity and flights to the odor source began for C. hemipterus on Day 4, but C. lugubris continued to walk to the source and rarely flew regardless of age. Both species displayed maximum response to food odors from 6 to 9 days after emergence and showed bimodality in host orientation during the photophase. C. hemipterus maintained with artificial diet took flight as frequently as when maintained with water only, as long as they were deprived of diet for 36 h before the test; however, after takeoff, beetles maintained with diet were less likely to fly to food odor in comparison to beetles maintained with water. C. lugubris maintained with diet until 36 h before the test displayed a reduction in walks upwind to sources of food odor compared to beetles maintained with water. Three hours of unrestricted locomotion under a high-pressure sodium lamp did not enhance upwind orientation to host odors by C. hemipterus maintained with diet or water, or by C. lugubris maintained with water; however, such preexposure to a sodium lamp resulted in reduced takeoffs in C. hemipterus maintained with artificial diet. Both species were attracted to all fruit and vegetable substrates offered; however, aseptic substrates were less attractive than were substrates inoculated with Saccharomyces cerevisiae Hansen or Candida krusei (Castellani) Berkhout for C. lugubris. Despite the broad host range exhibited by these two nitidulid species, each responded to chemical cues from longrange (2.5 m), a trait once assigned to specialists.     

Behavior of Carpophilus hemipterus in a vertical flight chamber: transition from phototactic to vegetative orientation

The freeflight behavior of Carpophilus hemipterus (L.) (Coleoptera: Nitidulidae) was investigated in a vertical flight chamber to measure takeoff propensity relative to beetle age and to determine the effect of food odors on phototactic orientation. A high-pressure sodium lamp presented from above induced a phototactic flight response, especially in beetles 3–7 days old. The majority of phototactic flights lasted less than 15 min for both males (73%) and females (85%), but ranged up to 100 min. A bimodal periodicity in flight propensity was recorded during the photophase with a small peak in activity occurring 14–10 h prior to scotophase and a large 4 h peak occurring from 3 h prior to scotophase to 1 h after when the onset of scotophase would normally have occurred. Beetles maintained with water as adults usually displayed higher levels of flight activity than did beetles maintained with artificial diet.After a period of vertical flight, photokinetic and phototactic response declined, and flight instability increased, as indicated by an overall decrease in the mean rate of climb, accompanied by an increase in the variability of this measure and an increase in horizontal displacement.When food odor (apple cider vinegar) was introduced the rate of climb dropped rapidly and beetles usually landed regardless of how long they had been in flight. When the food odor was removed, takeoff occurred and the beetle returned to its previous rate of climb. When food odor was repeatedly introduced during the same flight, there was no apparent cumulative effect, and each time it was removed, the beetle re-initiated phototactic flight. Unlike true migratory flight in which response to vegetative cues is temporarily inhibited until the insect has engaged in a period of phototactic flight, C. hemipterus flight could be better characterized as extended foraging in which phototactic flight is readily interrupted by encounters with food odor.

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Résumé Le comportement de vol libre de C. hemipterus L. a été examiné dans une chambre à vent verticale pour déterminer la tendance au vol après l'émergence et observer l'influence de l'odeur d'aliments sur le vol phototactique. Une lampe à haute pression de sodium induit une réponse de col phototactique, qui débute le 3^e jour et reste puissante jusqu'au 8^e jour. La majorité des vols phototactiques duraient moins de 15 pour les mâles (73%) et les femelles (85%), mais pouvait se prolonger jusqu'à 100. Les femelles nourries sur régime artificiel présentaient la plus forte proportion de vol dépassant 35. Cependant, les adultes maintenus exclusivement en présence d'eau ont présenté un niveau d'activité de vol encore supérieur. Une tendance au vol à périodicité quotidienne bimodale a été notée avec un petit pic d'activité à 14–10 heures avant la scotophase et un grand pic durant 4 heures, de 3 heures avant la scotophase à une heure après le début de la scotophase.Après une période de vols verticaux, les réactions photocinétiques et phototactiques ont diminué, et l'instabilité des vols augmenté comme en a témoigné la diminution du taux total d'ascensions et l'accroissement de la variabilité de ce paramètre et des déplacements horizontaux. L'introduction d'odeurs d'aliments pendant le vol phototactique provoque une diminution immédiate du taux d'ascension et conduit généralement à un atterrissage indépendamment du temps antérieur de vol. Quand l'odeur était supprimée le décollage avait lieu et l'insecte retrouvait sont taux antérieur d'ascension. Quand la même odeur d'aliment a été réintroduite à répétition pendant le même vol, il n'y a pas eu d'effets cumulatifs apparents. Par opposition avec d'autres insectes présentant un véritable comportement migrateur pour lesquels la désinhibition des réactions végétatives ne se produit qu'après une longue période de locomotion continue, le vol de C. hemipterus est mieux caractérisé comme un comportement de prospection pour lequel les aliments inhibent immédiatement la dispersion.

     

Pheromone-mediated mass trapping and population diversion as strategies for suppressing Carpophilus spp. (Coleoptera: Nitidulidae) in Australian stone fruit orchards

1 Five experiments were conducted during 1995–99 in stone fruit orchards on the Central Coast and in inland New South Wales, Australia, on the use of synthetic aggregation pheromones and a coattractant to suppress populations of the ripening fruit pests Carpophilus spp. (Coleoptera: Nitidulidae). 2 Perimeter‐based suppression traps baited with pheromone and coattractant placed at 3 m intervals around small fruit blocks, caught large numbers of Carpophilus spp. Very small populations of Carpophilus spp. occurred within blocks, and fruit damage was minimal. 3 Carpophilus spp. populations in stone fruit blocks 15–370 m from suppression traps were also small and non‐damaging, indicating a large zone of pheromone attractivity. 4 Pheromone/coattractant‐baited suppression traps appeared to divert Carpophilus spp. from nearby (130 m) ripening stone fruit. Ten metal drums containing decomposing fruit, baited with pheromone and treated with insecticide, attracted Carpophilus spp. and appeared to reduce populations and damage to ripening fruit at distances of 200–500 m. Populations and damage were significantly greater within 200 m of the drums and may have been caused by ineffective poisoning or poor quality/overcrowding of fruit resources in the drums. 5 Suppression of Carpophilus spp. populations using synthetic aggregation pheromones and a coattractant appears to be a realistic management option in stone fruit orchards. Pheromone‐mediated diversion of beetle populations from ripening fruit may be more practical than perimeter trapping, but more research is needed on the effective range of Carpophilus pheromones and the relative merits of trapping compared to attraction to insecticide‐treated areas.     

Comparison of trap designs for use with aggregation pheromone and synthetic co-attractant in a user-friendly attract and kill system to control Carpophilus spp. (Coleoptera: Nitidulidae)

Abstract  Three field experiments were conducted in stone fruit orchards in the Goulburn Valley, northern Victoria, Australia to identify a user-friendly trap for use in attract and kill stations for control of Carpophilus spp. (Coleoptera: Nitidulidae). A funnel trap design was compared with two types of delta trap in one experiment and two types of Lucitrap in another experiment. All traps were baited with synthetic co-attractant and synthetic pheromone. The funnel trap was the most effective type of trap tested. The funnel traps caught significantly more beetles than either the delta traps with bio-attractant or delta traps wrapped with insecticide impregnated banana wrap. No difference was observed between catches in either delta trap design. Lucitraps without covers caught significantly more beetles than Lucitraps with covers but funnel traps caught significantly more beetles than either type of Lucitrap. Beetle numbers caught in the funnel traps were three to five times higher than in Lucitrap. The use of funnel traps in attract and kill stations to protect stone fruit crops by suppressing the Carpophilus spp. population gave superior control to 'grower normal practice of spraying insecticides' both in terms of Carpophilus spp. numbers and in terms of reduction in percentage of fruit damaged.     

Autodissemination of Beauveria bassiana by Sap Beetles (Coleoptera: Nitidulidae) to Overwintering Sites

An autoinoculative device was used to test the ability of sap beetles (Coleoptera: Nitidulidae) to carry a specific strain of Beauveria bassiana (Balsamo) Vuillemin to overwintering sites in a multiyear field study. The device was baited with the pheromone and coattractants for the dusky sap beetle (Carpophilus lugubris Murray) and placed in the field in the fall of each year. The introduced strain occurred at high frequency among the B. bassiana isolated in the fall of all four years tested (100% of all isolates from 21 of 22 collection dates). The introduced strain of B. bassiana was isolated at high frequency from all the B. bassiana-contaminated sap beetles recovered from the overwintering traps (100% of all isolates from 13 of 23 trap/beetle species combinations) and was highest after the longest fall exposure. The introduced strain was primarily isolated from C. lugubris and C. antiquus, but species distribution was also dependent on the overwintering trap design used. Few non-sap beetles species of insects were recovered from the artificial overwintering sites. Although B. bassiana was isolated from free flying sap beetles caught in traps in the spring of each year, none were infected with the introduced strain. The autoinoculating device provides selective contamination of sap beetles in overwintering sites when used in the fall. It may be useful in providing some control of sap beetles or other insects where limited numbers of mass overwintering sites (such as tree holes) occur.     

Susceptibility of Sap Beetles (Coleoptera: Nitidulidae) to Entomopathogenic Nematodes

Nitidulid beetles (Coleoptera) are considered to be serious pests of date palms throughout the world. They attack ripe fruit, causing it to rot, and damage is reflected in both reduced yield and lower fruit quality. The present study was aimed at an evaluation of the susceptibility to different sap beetles to entomopathogenic nematodes. We further tested nematode efficacy in pots filled with soil infested by third instar larvae of the two beetle species. In Petri dish assay, mortality levels of Carpophilus humeralis and C. hemipterus exposed to Heterorhabditis sp. IS-5 strain indicated that the latter is less susceptible to nematode infection. Exposure of both sap beetle species to different nematode strains gave moderate levels of mortality (35-65%) with the heterorhabditid strains HP88, IS-5 and IS-25. The IS-12 strain of Heterorhabditis sp. showed poor virulence (<35% mortality) against larvae of C. humeralis as well as larvae and pupae of C. humipterus. The nematode species S. riobrave showed moderate virulence (35-65%) mortality to larvae and pupae of S. humeralis as well as to larvae of C. hemipterus . Exposure of C. hemeralis to different concentrations of Heterorhabditis sp. IS-5 in pots containing soil resulted in high mortality (>65%). In contrast, the lower concentrations (500 and 1000 nematodes/pot) caused low mortality (35%) of C. hemipterus . Other heterorhabditid strains caused 95-100% mortality of C. humeralis in pot assay. The HP88 strain of H. bacteriophora and the Tx strain of Steinernema riobrave showed poor effectiveness. Incubation of different nematode strains with the C. humeralis larvae at high temperature (32 C) resulted in an increase in insect mortality with the IS-12 and IS-21 strains. Reduced mortality was recorded with the HP88 strain treatment at the higher temperature. The IS-5 and IS-12 strains were equally effective in all three soil types tested, whereas the IS-19 strain was more effective in the Almog type soil than in the others.     

Responses of Carpophilus hemipterus larvae and adults to selected secondary metabolites of maize

Selected secondary metabolites produced by maize (Zea mays L.) were tested for effects on larvae and adults of the dried-fruit beetle [Carpophilus hemipterus (L.)] in no-choice and choice assays. Feeding by adults and larvae was significantly reduced by ferulic acid and 6-methoxy-2-benzoxazolinone (MBOA) in no-choice assays. In choice assays, larvae and adults generally preferred diets with trans-cinnamic acid, quercetin, rutin, and thymol, but were repelled by diets with either ferulic acid or MBOA.

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Résumé Les réactions de larves et adultes du nitidulidé C. hemipterus (L.), vecteur de champignons produisant la mycotoxine, aux composés phénoliques caractéristiques, aux flavonoïdes et aux acides hydroxamiques, métabolites secondaires qui provoquent la résistance du maïs (Zea mays L.) ont été examinées au cours d'expériences avec et sans choix. L'alimentation des adultes et des larves est généralement réduite par les acides coumarique et férulique et par la 6-méthoxy-2-benzoxazolinone dans des expériences sans choix; les insectes évitent généralement les aliments qui contiennent ces produits. Quoi qu'il en soit, les larves préfèrent consommer d'autres aliments contenant les autres phénoliques ou flavonoïdes examinés. Les adultes sont plus inconstants dans leur choix alimentaires, mais préfèrent souvent des aliments contenant de la quercetine. Ainsi, des programmes de sélection orientés contre les principaux ravageurs comme Heliothis zea (Boddie) ou Ostrinia nubilalis (Hübner), impliquant la sélection de plantes à teneur élevée en acides phénolique ou hydroxamique, augmentant probablement aussi la résistance aux nitidulidés.

     

A new species of Carpophilus Stephens (Coleoptera: Nitidulidae) from Florida,U.S.A., and a reappraisal of the type material of Carpophilus ophthalmicus Murray

A new species of Carpophilus is described from males and females taken in Florida, U.S.A. A lectotype male and paralectotype female are chosen for Carpophilus ophthalmicus, and a new species is described from material misidentified as ophthalmicus.     

Dusky Sap Beetle Mediated Dispersal of Bacillus subtilis to Inhibit Aspergillus flavus and Aflatoxin Production in Maize Zea mays L.

Laboratory assays were performed with detached milk stage maize ( Zea mays L.) ears and dusky sap beetles ( Carpophilus lugubris Murray) carrying the Kodiak Concentrate formulation of the bacterium, Bacillus subtilis (Ehrenberg) Cohn. After 1 day of exposure to the B. subtilis- contaminated C. lugubris , the colonization of mechanically damaged kernels by Aspergillus flavus Link ex. Fries was reduced from 82% (if the A. flavus was inoculated first) to 41% (if B. subtilis was added by C. lugubris before the A. flavus ). Field cage studies were performed with an autoinoculative device containing B. subtilis into which C. lugubris beetles were introduced. C. lugubris -dispersed B. subtilis reduced visible A. flavus colonization by 97% when the A. flavus was added to purposely damaged maize ears 4 days after C. lugubris were released from the autoinoculator. In 1993 field studies, none of the purposely damaged ears that allowed access to C. lugubris beetles emerging from autoinoculators containing B. subtilis had visible sporulating A. flavus compared with 92% of ears that did not allow access of C. lugubris but that subsequently had the A. flavus inoculum added. In 1994 field studies, 70% of the ears that excluded C. lugubris had aflatoxin levels greater than 200 ppb in purposely damaged kernels, as opposed to less than 10% of kernels that permitted access by natural populations of C. lugubris that probably acquired B. subtilis from a single autoinoculator. Aflatoxin levels in these ears were negatively correlated with the presence of both B. subtilis and C. lugubris . The B. subtilis was widely dispersed over a 16-ha area as indicated by maize ear and C. lugubris trap sampling. These studies indicate that autoinoculative dispersal of B. subtilis by natural populations of C. lugubris is a potentially useful means for reducing A. flavus and aflatoxin in maize.     

Molecular phylogenetics of a South Pacific sap beetle species complex (Carpophilus spp., Coleoptera: Nitidulidae)

Several species of sap beetles in the genus Carpophilus are minor pests of fresh produce and stored products, and are frequently intercepted in biosecurity operations. In the South Pacific region, the superficially similar species C. maculatus and C. oculatus are frequently encountered in these situations. Three subspecies of C. oculatus have been described, and the complex of these four taxa has led to inaccurate identification and questions regarding the validity of these taxa. A molecular phylogenetic study using the mitochondrial gene cytochrome c oxidase I (COI) and two nuclear markers comprising the rDNA internal transcribed spacer 2 (ITS2) and the D1-D2 region of the large (28S) ribosomal RNA subunit showed that C. maculatus, and C. o. cheesmani were easily differentiated from the two other subspecies of C. oculatus. COI also showed differentiation between C. o. gilloglyi and C. o. oculatus, but this was not shown when third codon positions were removed and when RY-coding analyses were conducted. Generalised mixed Yule-Coalescent (GMYC) models were fitted to trees estimated from the COI data and were analysed using a multimodel approach to consider the evidence for three taxonomic groupings of the C. oculatus group. While the arrangement with the highest cumulative weight was not the arrangement ultimately accepted, the accepted taxonomy also had an acceptable level of support. ITS2 showed structure within C. oculatus, however C. o. oculatus was resolved as paraphyletic with respect to C. o. gilloglyi. COI showed evidence of sequence saturation and did not adequately resolve higher relationships between species represented in the dataset. 28S resolved higher relationships, but did not perform well at the species level. This study supports the validity of C. maculatus as a separate species, and provides sufficient evidence to raise C. o. cheesmani to the level of species. This study also shows significant structure within and between C. o. gilloglyi and C. o. oculatus, giving an indication of recent speciation events occurring. To highlight the interesting biology between these two taxa, C. o. gilloglyi is retained as a subspecies of C. oculatus. These results give clarity regarding the taxonomic status of C. maculatus and the subspecies of C. oculatus and provide a platform for future systematic research on Carpophilus.     

The effect of antennectomy on the humidity reactions of Carpophilus dimidiatus and C. hemipterus (Coleoptera, Nitidulidae)

The humidity reactions of intact and antennaless specimens of Carpophilus dimidiatus and C. hemipterus were studied. The results suggested that the hygroreceptors mediating a response to high humidities are situated on the antennae in both species and those mediating a response to low humidities, in the case of C. dimidiatus, elsewhere on the body.

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Résumé On a étudié la réaction à l'humidité de Carpophilus hemipterus et C. dimidiatus intacts et sans antennes. Les résultats suggèrent que les hygrorécepteurs qui répondent aux humidités élevées se trouvent sur l'antenne dans les deux espèces et ceux qui répondent aux basses humidités sur l'antenne de C. hemipterus et dans une autre partie du corps de C. dimidiatus.

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On transfer from tropical Stored Products Centre, O.D.M., Slough, Bucks