Functional response and mutual interference in the parasitoid Coptera haywardi (Oglobin) (Hymenoptera: Diapriidae) attacking Anastrepha ludens (Loew) (Diptera: Tephritidae) pupae.

Functional response and mutual interference are important attributes of natural enemies that should be analysed in species with the potential to be used as biological control agents in order to increase the predictive power of the possible benefits and/or consequences of their release in the field. Our main objective was to determine the functional response and mutual interference of Coptera haywardi (Oglobin), a pupal parasitoid of economically important fruit flies (Diptera: Tephritidae). The functional response of C. haywardi on A. ludens pupae corresponded to a type II model, with an attack rate of 0.0134 host pupa/h and a handling time of 1.843 h, which reveals a meticulous selection process of pupal hosts. The effect of mutual interference among foraging females was negatively correlated with increased parasitoid density in the experimental arena, showing a gradual decline in attack rate per individual female. The increase in the number of foraging females also had an impact on the number of oviposition scars per pupa and the number of immature parasitoids per dissected pupa, but not on the percentage of adult emergence or the sex ratio. Our results suggest that C. haywardi could act as a complementary parasitoid in the control of fruit fly pupae, since the random distribution of these pupae in the soil would decrease the possibility of aggregation and mutual interference between foraging females.

Shadow Effect for Small Insect Detection by W-Band Pulsed Radar.

In radar entomology, one primary challenge is detecting small species (smaller than 5 cm) since these tiny insects reflect radiation that can be poorly observable and, therefore, difficult to interpret. After a literature search on radar entomology, this research found few works where it has been possible to sense insects with dimensions smaller than 5 cm using radars. This paper describes different methodologies to detect Mediterranean fruit flies with 5-6 mm sizes using a pulsed W-band radar and presents the experimental results that validate the procedures. The article's main contribution is the successful detection of Mediterranean fruit flies employing the shadow effect on the backscattered radar signal, achieving an 11% difference in received power when flies are present. So far, according to the information available and the literature search, this work is the first to detect small insects less than 1 cm long using a pulsed radar in W-Band. The results show that the proposed shadow effect is a viable alternative to the current sensors used in smart traps, as it allows not only detection but also counting the number of insects in the trap.

Bacterial Diversity Associated with Anastrepha obliqua Males Change Under Mass-Rearing Conditions and with Irradiation.

Recent advances in understanding the symbiotic interactions between bacteria and fruit flies have shown that they are relevant for mass rearing and the sterile insect technique (SIT). SIT involves mass production and release of sterile insects that would copulate with their wild conspecifics and thus decrease the population growth rate. The irradiation process used to sterilize mass-reared flies can modify the diversity and structure of the midgut bacterial communities, which could affect sterile male survival, flight capacity, and sexual competitiveness. Our aim was to compare bacterial communities in the midgut of wild and mass-reared Anastrepha obliqua (Macquart) males irradiated at 0, 60, and 80 Gy. After adult's emergence, their midguts were dissected, DNA was extracted, and high-throughput sequencing of the V3-V4 region of the 16S rDNA gene was performed. A total of 11 phyla, 17 classes, 47 families, and 52 genera of bacteria were identified. The most representative phylum was Proteobacteria and the predominant family was Enterobacteriaceae. We found that wild males had a different intestinal bacterial community from mass-reared males. In addition, irradiation at 60 and 80 Gy caused changes in the diversity and structure of the midgut microbiota of these sterile males, suggesting that mass rearing and irradiation cause artificial selection of the bacterial communities in the gut of A. obliqua males.

Superparasitism and fitness parameters in three native wasp parasitoids (Braconidae: Opiinae) of the Mexican fruit fly, Anastrepha ludens (Diptera: Tephritidae).

Knowledge on reproductive strategies and host use in insect parasitoids is fundamental for biological control purposes. Superparasitism and multiparasitism, oviposition in a previously parasitized host by a female of the same or different species, respectively, may impact pest management decisions. Our objective was to determine the occurrence of superparasitism and multiparasitism in three species of native larval-pupal solitary endoparasitoids that attack Anastrepha Shiner species (Diptera: Tephritidae) in the Neotropical region, and the possible effect on offspring fitness parameters. Doryctobracon crawfordi (Viereck), Utetes anastrephae (Viereck), and Opius hirtus (Fischer) occur in sympatry in Mexico, and are currently under consideration for use as biocontrol agents. Experiments were conducted under laboratory conditions with females acting alone (self-superparasitism), females in groups of the same species (conspecific superparasitism), and females in mixed groups (multiparasitism). Our results showed that self-superparasitism is an uncommon strategy in the three native species and is rare under conditions of intraspecific competition. In the case of multiparasitism, a higher number of immature stages of U. anastrephae was observed, compared to those of D. crawfordi and O. hirtus. However, it is not clear yet if this was due to some adult female trait or to the competitive ability of the larvae. We conclude that most females of the native species studied appeared to avoid superparasitism, specifically when acting alone, suggesting a high discrimination ability, which is probably a result of a close relationship and evolutionary history with Anastrepha hosts.

Does body size influence mating success? A morphometric study of two Anastrepha (Diptera: Tephritidae) fruit fly species.

Male mating success depends on various traits and factors, and correctly identifying these traits can be key in the context of pest management. For tephritid pests, controlled through the sterile insect technique (SIT) traits, such as male size, can be manipulated through mass-rearing procedures. Thus, it is particularly important to understand whether male size can favor mating success. Here, we evaluated mating success of males of different sizes in Anastrepha ludens and Anastrepha obliqua, two species controlled through SIT. For both species, a morphometric analysis was performed of mated and unmated mass-reared and wild males in field cages. In both A. ludens and A. obliqua, wild females did not discriminate their mates based on male size and mated more frequently with wild males regardless of size. For mass-reared males, we found no evidence of an advantage of the large males compared to small males in mating success. However, we did find differences between the morphometric traits of mass-reared and wild males. In A. ludens, traits associated to mating success were Face Width (FW), Head Width (HW), Thorax Length (TL) and Wing Length (WL), and for A. obliqua were FW, HW, WL and WW (Wing Width). Overall, FW and TL were more consistent predictors of mating success. In conclusion, female choice seems to suggest multivariate selection, confirming that overall body size (expressed as pupal diameter, which is highly correlated with weight) is not a decisive factor in male mating success in these two species. However, morphological traits such as FW, HW, TL, WL and WW may be relevant in mating preference of wild female.

Interaction Between Beauveria bassiana (Hypocreales: Cordycipitaceae) and Coptera haywardi (Hymenoptera: Diapriidae) for the Management of Anastrepha obliqua (Diptera: Tephritidae).

The interaction between the entomopathogenic fungus Beauveria bassiana (Balsamo) and the parasitoid Coptera haywardi (Oglobin), as potential biological control agents for Anastrepha obliqua (Macquart) fruit flies, was evaluated under laboratory and semi-protected field cage conditions. The effects of the parasitoids and fungus were individually and jointly assessed in Plexiglas cages. Application of B. bassiana dry conidia to soil produced 40% mortality in A. obliqua adults. However, mortality was lower (21.2%) on evaluation under field cage conditions. According to the multiple decrement life table analysis, the probability of death of A. obliqua was 88% when C. haywardi parasitoids and B. bassiana conidia were used in conjunction, 89% when only C. haywardi parasitoids were released and 23% when only B. bassiana conidia were applied. These results demonstrate that no synergistic, additive or antagonistic interaction took place with the simultaneous use of these natural enemies, since the presence of B. bassiana had no effect on the C. haywardi parasitism. These results indicate that the parasitoid is a better natural enemy for the control of A. obliqua, and show that, although the two biological control agents can be used simultaneously, their joint application will not produce increased control.

Variation in the cuticular hydrocarbons of the Mexican fruit fly Anastrepha ludens males between strains and age classes.

In this study cuticular hydrocarbons (CHCs) were characterized from wings of individual unmated males of different Anastrepha ludens (Loew) mass-reared strains of different ages (3 and 19-day-old): (a) a standard mass-reared colony (control), (b) a genetic sexing strain, (c) a selected strain, (d) a hybrid strain, and (e) wild males. We found that the hydrocarbon profiles in all males included two n-alkanes, five monomethyl alkanes, and two alkenes. CHCs ranged from C26 to C31 . The most prominent peaks were 2-methyloctacosane (2-Me-C28), n-nonacosene (C29:1), 2-methyltriacontane (2-Me-C30), and n-hentriacontene (C31:1). Significant variations in the CHC amounts of the mass-reared strains were observed from Day 9 and thereafter. Comparison of CHCs using multivariate and canonical analyses across ages and among mass-reared strains and wild males revealed qualitative and quantitative differences. The relative amounts of C29:1 and 2-Me-C30 were significantly higher across age groups in the mass-reared strains than those in the wild males. In contrast, amounts of n-nonacosane (C29) significantly increased in wild males as they aged. Through statistical analyses, we inferred that CHC amounts vary with age. Wild males differed significantly from the mass-reared strains in the amount of C29, and the genetic sexing strain Tap-7 had significantly higher values for 2-methylhexacosane (2-Me-C26). In contrast the selected and control strain differed from the other strains in amounts of C29:1 and 2-Me-C30. We suggest that differential profiles in hydrocarbon composition among the strains may be mainly due to environmental pressures.

The Effect of Application Rate of GF-120 (Spinosad) and Malathion on the Mortality of Apis mellifera (Hymenoptera: Apidae) Foragers.

Beneficial organisms like the honey bee, Apis mellifera L. (Hymenoptera: Apidae), are heavily affected by pest control practices that incorporate insecticides. Safer alternatives as the spinosad-based formulation GF-120 have been developed to overcome this issue. Though both the low concentration of spinosad and the ultra-low-volume application rate of GF-120 are supposed to have a low acute toxicity in honey bee foragers, to our knowledge such claims have not been explicitly proven. We thus carried out a series of experiments to assess the effect of GF-120, malathion, and Spintor (spinosad) on honey bee foragers when applied at two concentrations (80 and 1,500 ppm) and two application rates (low density rate [LDR]­80 drops of 5 mm diameter per square meter; high density rate [HDR]­thousands of 200 -µm-diameter droplets per square meter). Interestingly, the three pesticides caused low mortality on foragers when applied at LDR-80, LDR-1,500, or HDR-80. However, HDR-1,500 caused a very high mortality. Based upon these results, we developed a computer program to estimate the average number of foragers that are exposed at LDR and HDR. We found that more foragers receive a lethal dose when exposed at HDR than at the other rates. Our results support the hypothesis that the impact of GF-120 and malathion upon honey bees is minimal when applied at LDR and that computer simulation can help greatly in understanding the effects of pesticides upon nontarget species.

Influence of Methoprene on Pheromone Emission and Sexual Maturation of Anastrepha obliqua (Diptera: Tephritidae) males.

It has been demonstrated that the application of juvenile hormone analog, methoprene, reduces the time required for sexual maturation and enhances mating success in several species of tephritid fruit flies. This study examined the effect of different concentrations of methoprene incorporated into the diet of adult flies and distinct sugar:protein (S:P) ratios on sexual maturity and pheromone emission of Anastrepha obliqua males. Diets with 0.2 and 0.5% of methoprene accelerated sexual maturation of males compared with untreated males. In subsequent assays, the enhancement of male pheromone emission and sexual maturation by the incorporation of 0.02% methoprene into a 24:1 (S: P) diet was confirmed. Among the volatiles released by males, (Z)-3-nonenol and (Z,Z)-3,6-nonadienol were emitted at higher quantities by flies treated with methoprene than untreated ones. The results show that methoprene accelerates sexual maturation of mass-reared A. obliqua males and increases their mating propensity. This would reduce the time required to attain sexual maturation by sterile males, thus decreasing fly handling costs and improving the efficacy of the sterile insect technique.

Escape of Mexican Fruit Flies from Traps Baited with CeraTrap and Effect of Lure Feeding on Reproduction and Survival.

McPhail type traps are commonly used to catch tephritid pests using liquid baits as attractants. Unfortunately, these traps allow that some flies escape after feeding on the bait. Recently, a long-lasting and nontoxic enzymatic-hydrolyzed protein, known commercially as CeraTrap, has been introduced as a novel liquid bait. Because feeding on hydrolyzed proteins can improve reproduction in many tephritids, questions remain as to whether CeraTrap could provide a nutritious meal to flies that ingest the bait but escape the trap. In this study, we analyzed the efficacy of CeraTrap baited traps in capturing Anastrepha ludens flies of two different physiological states and two different diets. We also explored the effect of feeding on CeraTrap on the mating performance of males, ovarian development and fecundity of females and fly survival. Results demonstrated that CeraTrap attracted more protein-deprived and immature flies of both sexes and 2-30% of flies that entered the trap managed to escape. However, after feeding on the bait for 24 h, males exhibited lower mating success, longer latency to mate and longer copula durations than males fed on either sugar or sugar+ protein. Interestingly, females fed CeraTrap for 24 h developed ovaries as large as those females fed on sugar+ protein, however, they laid no eggs. When feeding continuously on CeraTrap, flies exhibited shorter lifespans and females laid no eggs. These findings indicate that, although some flies can ingest the bait and escape from the trap, their reproductive traits are negatively affected.

Behavioral response of two species of stingless bees and the honey bee (Hymenoptera: Apidae) to GF-120.

We present the results of evaluating the response of three species of bees, Trigona fulviventris (Guérin), Scaptotrigona mexicana (Guérin-Meneville), and Apis mellifera (L.), to food sources baited with the toxic bait GF-120 (NF Naturalyte), a spinosad-based bait exclusively used to manage fruit flies. Groups of foragers were trained to collect honey and water from a feeder located 50 m from the colonies. Once a sufficient number of foragers were observed at the experimental location, the training feeder was changed to two or three feeders that offered either honey and water, GF-120, Captor (hydrolyzed protein), GF-120 and honey (4:6), or Captor and honey (1:19). T fulviventris and S. mexicana rarely visited GF-120, Captor, or their mixtures with honey, while approximately 28.5 and 1.5% of A. mellifera foragers visited the GF-120 and honey and Captor and honey mixtures, respectively. Our results show that GF-120 clearly repels T. fulviventris and S. mexicana, whereas for A. mellifera, repellence is not as marked when GF-120 is combined with highly nutritious substances like honey.

Estimation of populations and sterility induction in Anastrepha luden (Diptera: Tephritidae) fruit flies.

The relationship between different release densities of sterile flies and fly trap captures, expressed as flies per trap per day, in the monitoring of Anastrepha ludens (Loew) populations was evaluated in mango orchards. The induction of sterility in fertile females was evaluated using different ratios of sterile: fertile males under field cage conditions. A direct relationship between recaptured flies and densities of release sterile flies was found. However, trap efficiency, expressed as percentage of recaptured flies, decreased as the density of released flies increased. Sterility induction was positively correlated to the ratio of sterile: fertile flies. A significant difference in egg fertility among treatments was observed. The trajectory of sterility induction slowed down after a sterile: wild ratio of 30:1, which suggests that this ratio could be appropriate in an sterile insect technique program with A. ludens. Sterility induction was greater when only sterile males were released than when releasing both sterile males and females, but the differences were not significant. Our findings contribute to a better interpretation of fly captures obtained from the field trapping networks, and to an improvement in the efficiency of sterile insect technique against A. ludens fruit flies, through the implementation of more rational sterile fly release densities.

Mass-Rearing Conditions Do Not Always Reduce Genetic Diversity: The Case of the Mexican Fruit Fly, Anastrepha ludens (Diptera: Tephritidae).

The application of the sterile insect technique (SIT) requires the adaptation of insects to mass-rearing conditions. It is generally accepted that this adaptation may include a reduction in genetic diversity and an associated loss of desirable characteristics for the effective performance of sterile insects in the field. Here, we compare the genetic diversity of two mass-reared strains of the Mexican fruit fly, Anastrepha ludens, and a wild (WIL) population collected near Tapachula, Mexico, using seven DNA microsatellites as molecular genetic markers. The mass-reared strains were a bisexual laboratory strain (LAB) with approximately 130 generations under mass-rearing and a genetic sexing strain, Tapachula-7 (TA7), also under mass-rearing for 100 generations. Our results revealed an overall low level of genetic differentiation (approximately 15%) among the three strains, with the LAB and WIL populations being genetically most similar and TA7 most genetically differentiated. Although there were some differences in allele frequencies between strains, our results show that overall, the adaptation to mass-rearing conditions did not reduce genetic variability compared to the wild sample in terms of heterozygosity or allelic richness, nor did it appear to alter the level of inbreeding with respect to the wild populations. These results are contrary to the general idea that mass-rearing always results in a reduction in genetic diversity. Overall, our findings can contribute to a better understanding of the impact that adaptation to mass-rearing conditions may have on the genetic make-up of strains.

Effect of the Sterile Insect Technique and Augmentative Parasitoid Releases in a Fruit Fly Suppression Program in Mango-Producing Areas of Southeast Mexico.

The Sterile Insect Technique (SIT), by means of sterile male releases of Anastrepha ludens (Loew), coupled with Augmentative Biological Control (ABC), by releasing the parasitoid Diachasmimorpha longicaudata (Ashmead), was evaluated in a commercial mango production area for one year. The obtained results were compared with mean fruit fly population values from two previous years without the combined use of both techniques. The treatments were: SIT + ABC, SIT, ABC, and Control, and each treatment was established in blocks of 5000 Ha separated by distances of 5-10 km. The evaluations were carried out through fruit sampling to assess percent parasitism and trapping of adult flies to obtain Flies per Trap per Day (FTD) values. The mean percentage of parasitism increased from 0.59% in the control treatment to 19.38% in the block with ABC. The FTD values decreased from ~0.129 and ~0.012 in the control block to 0.0021 in the block with SIT and ABC, representing a 98% suppression. The difference between the two periods in the control block was not significant. We conclude that the integration of both techniques resulted in an additive suppression of the pest population, supporting the use of both control techniques in an area-wide pest management context.

Release of sterile Aedes aegypti mosquitoes: chilling effect on mass-reared males survival and escape ability and on irradiated males sexual competitiveness.

In the sterile insect technique, it is important to measure the impact of mass-rearing and handling of sterile males to allow a successful control of the target wild population. This study evaluates the effect of pre-release chilling on the survival, escape ability, and sexual competitiveness of male Aedes aegypti. To determine survival and escape ability, mosquitoes were chilled at 4 °C using four different treatments of either one exposure (25 min) or two consecutive exposures (25 + 25 min, 25 + 50 min, 25 + 100 min). For sexual competitiveness, two different treatments were evaluated, chilling for 25 min once and twice. Results showed that the longest exposure to chilling caused a significant reduction of survival time, from 67 to 54 days. Escape ability was reduced by the first chilling from 25 to 7% and with the second chilling, it was reduced from 30 to 24% in the control to 4.9, 2.0 and 0.5% for 25, 50 and 100 min, respectively. Sexual competitiveness index was reduced from 1.16 in the control, to 0.32 and - 0.11 for treatments involving one and two chilling periods, respectively. It is recommended to increase the chilling temperature and reduce the exposure time to reduce the harmful effects on sterile males.

Can Entomopathogenic Nematodes and Their Symbiotic Bacteria Suppress Fruit Fly Pests? A Review.

Fruit flies (Diptera: Tephritidae) are serious pests that affect fruit production and marketing. Both third instar larvae and pupae are biological stages that persist in the soil until adult emergence. Entomopathogenic nematodes (ENs) are biological control agents that are used to control agricultural pests in greenhouse or field conditions. Several studies have been carried out under laboratory and field conditions showing how ENs can be applied within an area-wide integrated pest management approach to control fruit fly species in orchards and backyard fruit trees. In this review, we analyze how soil physical characteristics and biotic factors affect the performance of these biological control agents. Of the reviewed papers, more than half evaluated the influence of soil texture, humidity, temperature, and other factors on the performance of infective juveniles (IJs). Abiotic factors that significantly influence the performance of IJs are temperature, humidity, and texture. Among the biotic factors that affect IJs are fungi, bacteria, mites, insects, and earthworms. We conclude that ENs have the potential to be applied in the drip area of fruit trees that are infested by fruit flies and contribute to their suppression. This approach, in conjunction with an area-wide pest management approach, may contribute to pest suppression and increase the sustainability of agroecosystems.

Immature stages of Utetes anastrephae (Hymenoptera: Braconidae) developed in Anastrepha fruit fly larvae (Diptera: Tephritidae).

The morphology of the immature stages of Utetes anastrephae (Hymenoptera: Braconidae), a native parasitoid of larvae of flies of the Neotropical genus Anastrepha (Diptera: Tephritidae), is shown. This study aimed to characterize the immature stages and morphological changes in the development of the koinobiont endoparasitoid in two species of larval hosts, Anastrepha obliqua and Anastrepha ludens. The definition of structures and morphological changes during development was made through daily microscopic observations and photographs of dissected hosts. The immature development of the parasitoid corresponds to a holometabolous insect with three well-defined stages: egg (two days), larva with three larval instars (approximately eight days), and pupa (six days). Similar development times were obtained in the two host species. Males and females completed their cycle in 17 and 18 days, respectively. During egg-first instar development, host antagonistic activity through melanization and encapsulation as mortality factors was evident and frequent only in A. obliqua. These results serve as basic knowledge for the use of this parasitoid in the biological control of fruit flies.

Survey of heritable endosymbionts in southern Mexico populations of the fruit fly species Anastrepha striata and A. ludens.

Heritable endosymbiotic bacteria associated with insects are ubiquitous and taxonomically diverse. Many of these endosymbionts influence the fitness of their hosts and/or manipulate their host reproduction. Exploiting the effects of endosymbionts on hosts for pest control is a growing research area, but requires knowledge of endosymbionts associated with the target pest population. In this study, we used molecular methods to screen southern Mexico populations of two species of tephritid fruit fly pests, Anastrepha ludens and A. striata, for heritable bacteria. The only heritable endosymbiont found was Wolbachia in A. striata. Based on multilocus sequence typing and phylogenetic analyses, this Wolbachia strain is new and belongs to the Wolbachia supergroup B. Wolbachia strains previously reported in members of the genus Anastrepha in South America belong to supergroup A. We discuss the potential implications for pest control of the presence of a different Wolbachia strain in southern Mexico.

Effect of a Probiotic-Enriched Diet on Sexual Competitiveness, Pheromone Emission, and Cuticular Hydrocarbons of Sterile and Fertile Anastrepha ludens (Diptera: Tephritidae).

The sterile insect technique has been used for the eradication or control of numerous tephritid fruit flies. However, mass-rearing and sterilization can affect the microbiota and sexual performance of male tephritid fruit flies. Despite the addition of postteneral protein food which contributes to the enhancement of the sexual performance of mass-reared males, in some cases, they are less competitive than their wild counterparts. Alternatively, the addition of probiotics may improve the sexual performance of mass-reared sterile males. In this study, we evaluated the effect of a postteneral Lactobacillus casei-enriched diet on the sexual competitivity, pheromone emission, and cuticular hydrocarbons of mass-reared sterile and fertile Anastrepha ludens (Loew) (Diptera: Tephritidae) males. Flies were fed either with sugar, standard diet (sugar and protein, 3:1), sugar + probiotic, or standard diet + probiotic. The addition of the probiotic improved the sexual competitivity of fertile and sterile males that were devoid of protein but led to a negative effect on males fed with a standard diet. As compared to males that were fed with the standard diet + probiotic/only sugar, the males fed with the standard diet or those fed on sugar + probiotic displayed a higher number of mating instances. Sterile males that fed on sugar + probiotic had a higher relative amount of anastrephine, epianastrephine, n-methyl octacosane, and 2-methyl triacontane than those fed on sugar only. Overall, these compounds were common in the treatments where males had the best sexual performance. Our results suggest that the probiotics offer nutritional advantages to males whose food lacks protein.

Effect of Aging on Three Lures Used for Monitoring Ceratitis capitata (Diptera: Tephritidae): Release Rate, Volatile Composition, and Fly Recaptures.

Prevention and control programs for Ceratitis capitata require a large supply of lures and traps for use in established trapping networks and mass-trapping suppression measures. The main lures currently used are: Trimedure (TML), three-component Biolure (BL), and Ceratrap (CT). The aim of this study was to determine the release rates of these lures, the chemical composition of their volatiles, and how these parameters change with exposure time. Tests were conducted under field conditions at three different elevations (25, 500, and 1,300 masl) during the dry and rainy seasons in Chiapas, Mexico. We found that for TML and BL, the release rate was similar in both seasons and at all three elevations. In the case of CT, the release rate was greater during the dry season and at the lowest elevation during the rainy season. With the caveat of using solid-phase microextraction technique for identification of lure compounds in this study, we found that the volatile compounds of TML were maintained throughout the rainy season, however, in the dry season, some compounds could not be detected. The volatile compounds emitted by BL were trimethylamine, ammonium acetate, and acetamide. Among volatile compounds of CT, acetic acid was the most abundant in the rainy season, while minor compounds were only detected during the first five weeks. Recapture rates were affected by elevation in the three lures tested and there was a significant interaction between elevation in exposure time for TML and BL.