Resistance to adult banded cucumber beetle (Coleoptera: Chrysomelidae) in romaine lettuce

Four cultivars of lettuce, Lactuca sativa L., were evaluated for their resistance to the adult banded cucumber beetle, Diabrotica balteata LeConte, under laboratory conditions. When paired with each of the other three cultivars, leaf consumption in all possible combinations of short-term (48 h) two-choice tests among the cultivars was significantly reduced only for 'Valmaine'. However, in a 48-h no-choice situation, beetles fed Valmaine or 'Short Guzmaine' had similarly low leaf consumption, followed by 'Parris White', with the greatest consumption occurring on 'Tall Guzmaine'. In longer term experiments, female beetles fed Valmaine for 10, 13, or 16 d generally had lower survival and the lowest body weights compared with beetles fed any of the other three cultivars. No mature eggs were found in the ovaries of females fed Valmaine, whereas from 14% (Short Guzmaine, day 10) to 100% (Tall Guzmaine, day 13) of females fed the other cultivars produced mature eggs. In a starvation test, most D. balteata of either sex did not survive after 7 d with access only to water. Moreover, starved females did not produce mature eggs. Thus, food consumption by adult D. balteata is very important to their survival and reproductive performance, and it is likely that females fed Valmaine failed to produce mature eggs because they did not consume a sufficient amount of this cultivar. However, because Valmaine-fed beetles maintained their body weight and lived significantly longer than starved beetles, it appears that they can obtain some nourishment from their limited feeding on this cultivar. Overall, these results suggest that Valmaine, and to a lesser extent short Guzmaine (a cultivar produced by crossing Valmaine with two other cultivars), exhibit antixenosis-based resistance against D. balteata.     

Banded cucumber beetle (Coleoptera: Chrysomelidae) resistance in romaine lettuce: understanding latex chemistry

Many plants subjected to herbivore damage exude latex, a rich source of biochemicals, which plays important roles in host plant resistance. Our previous studies showed that fresh latex from Valmaine, a resistant cultivar of romaine lettuce Lactuca sativa L., applied to artificial diet is highly deterrent to feeding by banded cucumber beetle, Diabrotica balteata LeConte, compared to the latex of a closely related susceptible cultivar Tall Guzmaine. The deterrent factor(s) could be extracted from Valmaine latex with water–methanol (20:80). In this study, further fractionation of the methanolic crude extract of Valmaine latex was performed using reverse-phase and ion-exchange solid-phase extraction to isolate the deterrent compounds. Retention of deterrent compounds on anion and cation exchange resin suggested the presence of highly polar compounds with both carboxylic and amine groups in Valmaine latex. Further bioassay-directed fractionation of cation exchange extract using LC/MS indicated the presence of at least 3 major and an unknown number of minor compounds in the bioactive fraction between 3 and 4 min. The m/z 210 out of the 3 major compounds showed strong amino acid characteristics (glutamine and/or glutamic acid) when subjected to further MS ⁿ degradation. Our studies suggest that nitrogenous ingredients of latex play a key role in Valmaine resistance to D. balteata, and latex may be a source of bioactive compounds with a potential use in pest management.     

Field and vegetable crops as hosts of larval western spotted cucumber beetle (Coleoptera: Chrysomelidae)

The western spotted cucumber beetle, Diabrotica undecimpunctata undecimpunctata Mannerheim, is an important pest of melons (Cucurmis melo L.) in northern California. Recent observations indicate that adults are using alfalfa (Medicago sativa L.) as a feeding host and larvae may be developing on the roots. Greenhouse studies were conducted during the winters of 2009 and 2010 in which larval development was compared on the roots of six field and vegetable crops commonly grown in the southern Sacramento Valley. The growth parameters used to evaluate the hosts were larval and pupal head capsule width, body width, and body length as well as total survival percentage and survival percentage to the third instar. According to larval growth and survivorship in 2009, maize (Zea mays L.) was the best host, followed by alfalfa and tomato (Solanum lycopersicum L.) that were roughly equivalent to one another. Melon was a slightly weaker host than alfalfa and tomato; sunflower (Helianthus annuus L.) was a completely incompatible host and thus dropped from the 2010 study. In 2010, melon was the weakest host for larval development. Maize was the superior host again followed closely by alfalfa that performed slightly better than tomato and lima bean (Phaseolus lunatus L.). Data suggest that larval western spotted cucumber beetles may primarily develop outside of melon fields and the adults emigrate to melons.     

Buttercup squash provides a marketable alternative to blue hubbard as a trap crop for control of striped cucumber beetles (Coleoptera: Chrysomelidae)

Winter squash is a vital agricultural commodity worldwide. In the Northeastern United States, the primary insect pest is the striped cucumber beetle, Acalymma vittatum F. Using a Blue Hubbard squash (Cucurbita maxima Duchesne) perimeter trap crop system can reduce insecticide use by >90% in butternut squash (C. moschata Poir), the primary winter squash grown in this region. Despite the savings in insecticide costs, growers may be reluctant to give up field space for a perimeter crop of Blue Hubbard squash, which comprises only 5% of the winter squash market in New England as compared with 19% for buttercup squash. Finding a more marketable trap crop would lower the barrier for adoption of this system. We tested eight varieties of three species of cucurbits for attractiveness to beetles relative to Blue Hubbard and butternut squash, and chose buttercup squash as the most promising replacement. We compared the effect of a buttercup border, Blue Hubbard border, or control (no border) on beetle numbers, herbivory, insecticide use, pollination, and pollen limitation in the main crop. We found that buttercup squash performed equally well as Blue Hubbard as a trap crop, with 97% reduction in total insecticide use compared with control fields. Honey bees (Apis mellifera L.) and squash bees (Peponapis pruinosa Say) were the predominant pollinators, and border treatments did not affect visitation. Hand pollination did not increase reproduction or yield, indicating that natural pollination was sufficient for full yield. This study confirms the effectiveness of perimeter trap crop systems and offers growers a more marketable trap crop for managing cucumber beetle damage.     

Male traits under cryptic female choice in the spotted cucumber beetle (Coleoptera: Chrysomelidae)

Males of the spotted cucumber beetle (Diabrotica undecimpunctatahowardi) rhythmically stroke females with their antennae duringcopulation. Males that stroke quickly have a higher probabilityof being accepted as a mate. We determined (1) the mechanismby which females prevent unattractive males from passing spermatophores,(2) whether antennal stroking signals to females the likelihoodof receiving a nuptial gift, and (3) if other male traits inaddition to stroking are subjected to sexual selection fromfemale preference. Dissections of pairs flash-frozen in copuladuring and after antennal stroking showed musculature that,when contracted, folded the vaginal duct leading to the female'sbursa copulatrix in a way that prevented complete penetrationby the aedeagus. These muscles were always contracted whilemales were stroking and always relaxed after stroking had ceased.Males accepted as mates did not differ from males that failedto pass a spermatophore in either absolute or relative bodyweight, aedeagus length, or the amount of cucurbitacins (potentialnuptial gifts) sequestered in their spermatophores. Although99% of the beetles that came to cucurbitacin-rich Cucurbitafruits in the field were males, males that had sequesteredcucurbitacins did not stroke females faster than males withno cucurbitacins, and fast-stroking males were not more likelyto find and sequester cucurbitacins than were males that strokedmore slowly. Males with a cucurbitacin slurry painted on theirantennae had no mating advantage over controls. We concludethat females discriminate among males after copulation hasbegun on the basis of antennal stroking displays (or some traitcorrelated with stroking speed) that males perform to enticefemales to relax their bursal sphincter.     

Yield response of pumpkin and winter squash to simulated cucumber beetle (Coleoptera: Chrysomelidae) feeding injury

Field studies were conducted to characterize the yield responses of pumpkin, Cucurbita pepo L., and wintersquash, Cucurbita maxima Duch., to simulated striped cucumber beetle, Acalymma vittatum (F.), feeding damage. Preliminary studies in pumpkin ('Spookie') were conducted in 1993, and more complete studies in 1994 and 1995 in pumpkin (Spookie), and in 1995 and 1996 in winter squash ('Waltham'). Plants were artificially injured 1 time at the cotyledon, 1st, 2nd, or 3rd true leaf stages by clipping specified percentages of leaf area from all leaves on the plant. Treatments consisted of a control treatment with no simulated injury and 4 levels of simulated injury. Pumpkin treatments were 10, 20, 40, and 80% leaf area removal in 1993 and 1994 and 20, 40, 60, and 80% in 1995. Winter squash treatments were 20, 40, 60, and 80% leaf area removal in 1995, and 10, 20, 40, and 80% in 1996. In 1993, 5 wk after planting, mean leaf, stem, and total dry weight were significantly lower on pumpkin plants subjected to > or = 20% simulated leaf injury regardless of the growth stage when the injury occurred. Injury at the 3rd leaf stage resulted in a significant reduction in the mean leaf and total dry weight of plant, across all levels of injury. At 2-3 wk before crop maturity, fruit weight and number of fruit per plant were significantly lower with 80% leaf area removal. In 1994 and 1995, and when pumpkins were grown to full maturity and apparently able to compensate for the simulated damage, neither number nor weight of marketable fruit per plant were affected significantly by simulated injury at any stage of development nor level of simulated injury. In contrast, > or = 20% and 80% leaf area removal to winter squash resulted in significant reductions in the weight of marketable fruit and/or number of marketable fruit, in 1995 and 1996, respectively. The results of this study indicate that pumpkins can tolerate relatively high levels of simulated defoliation, whereas winter squash is less tolerant. Variable results across years in winter squash may indicate an interaction between injury and plant stress caused by abiotic factors such as soil moisture availability.     

Feeding behavior of leaf beetles (Coleoptera,Chrysomelidae)

The feeding behavior of adult leaf beetles (41 species from 18 genera and 8 subfamilies) was studied for the first time. Beetles of the genera Chrysolina, Chrysomela, Cryptocephalus, Galeruca, Gastrophysa, Labidostomis, Leptinotarsa, Timarcha, and Cassida stigmatica gnaw a leaf from the edge, whereas the representatives of Donacia, Galerucella, Lema, Lilioceris, Oulema, Phyllobrotica, Plagiodera, Zeugophora, Hypocassida, and most species of Cassida gnaw the leaf plane. In addition, adults of Lilioceris merdigera and Donacia clavipes feed on young leaves rolled into a tube. New host plants are reported for the first time: Hyoscyamus niger for the larvae of the Colorado potato beetle and Naumburgia thyrsiflora for Galerucella grisescens.     

Anthophagy in the leaf beetles (Coleoptera,Chrysomelidae)

Laboratory experiments were performed on leaf beetles collected on flowers. Flower consumption (anthophagy) was demonstrated for adults of Donacia bicolora, D. brevitarsis, D. obscura, D. thalassina, Plateumaris discolor, Cryptocephalus laetus, C. sericeus, C. solivagus, Labidostomis longimana, Hydrothassa marginella, Phaedon concinnus, Galerucella nymphaeae, Neocrepidodera femorata, Altica oleracea, Aphthona lutescens, Longitarsus pellucidus, and larvae of Entomoscelis adonidis, H. marginella, and G. nymphaeae for the first time for all the species except for P. discolor, L. longimana, and C. sericeus. The feeding behavior was described, and the mouthparts of adults were studied. The mandibles and maxillae of the Donacia, Plateumaris, and Orsodacne species mentioned are adapted to consuming pollen. Some questions of ecology and evolution of anthophagy are discussed. An overview of literature on anthophagy in the leaf beetles is presented.     

Potential resistance of crape myrtle cultivars to flea beetle (Coleoptera: Chrysomelidae) and Japanese beetle (Coleoptera: Scarabaeidae) damage

Field and laboratory studies were conducted to identify potential resistance among crape myrtles, Lagerstroemia spp., to Japanese beetle, Popillia japonica Newman and to flea beetles, Altica spp. Damage ratings revealed variation among cultivars in susceptibility to beetle feeding. Cultivars with Lagerstroemia fauriei Koehne in their parentage exhibited the least amount of damage in choice and no-choice experiments, with few exceptions. The data indicate that both beetle species cause more feeding damage on certain cultivars of Lagerstroemia indica L., such as 'Country Red', 'Twilight', and 'Carolina Beauty' than interspecific cultivars with L. fauriei in their parentage, such as 'Natchez', 'Tonto', and 'Muskogee'. When comparing the effect of parentage on all of the major pests of crape myrtle, L. faurei confers resistance to all pests except crape myrtle aphid. No correlation was found between leaf toughness, leaf color, and leaf nutrients in estimating flea beetle cultivar preference. With this information, growers can more effectively target scouting measures to the most susceptible cultivars. and breeders can select plants that will require the fewest chemical inputs.     

Molecular phylogeny of rootworms and related galerucine beetles (Coleoptera: Chrysomelidae)

The Galerucinae (Coleoptera: Chrysomelidae) sensu stricto (true galerucines) comprise a large assemblage of diverse phytophagous beetles containing over 5000 described species. Together with their sister taxon, the flea beetles, which differ from true galerucines by having the hind femora usually modified for jumping, the Galerucinae sensu lato comprises over 13 000 described species and is the largest natural group within the Chrysomelidae. Unlike the flea beetles, for which robust hierarchical classification schemes have not been erected, an existing taxonomic structure exists for the true galerucines, based mostly on the works of the late John Wilcox. In the most recent taxonomic list of the Galerucinae sensu stricto, five tribes were established comprising 29 sections housing 488 genera. The majority of the diversity within these tribes is found within the tribe Luperini, in which two genera, Monolepta and Diabrotica, are known to contain over 500 described species. Here, we extend the work from previous phylogenetic studies of the Galerucinae by analysing four amplicons from three gene regions (18S and 28S rRNA; COI) representing 249 taxa, providing the largest phylogenetic analysis of this taxon to date. Using two seven‐state RNA models, we combine five maximum likelihood models (RNA + DNA for the rRNAs; three separate DNA models for the COI codon positions) for these partitions and analyse the data under likelihood using Bayesian inference. The results of these two analyses are compared with those from equally weighted parsimony. Instead of choosing the results from one optimality criterion over another, either based on statistical support, tree topology or philosophical predisposition, we elect to draw attention to the similar results produced by all three analyses, illustrating the robustness of the data to these different analytical methods. In general, the results from all three analyses are consistent with each other and previous molecular phylogenetic reconstructions for Galerucinae, except that increased taxon sampling for several groups, namely the tribes Hylaspini and Oidini, has improved the phylogenetic position of these taxa. As with previous analyses, under‐sampled taxa, such as the Old World Metacyclini and all sections of the subtribe Luperina, continue to be unstable, with the few taxa representing these groups fluctuating in their positions based on the implemented optimality criterion. Nonetheless, we report here the most comprehensive phylogenetic estimation for the Galerucinae to date.     

Spatial distribution and sampling plans for Mesoplatys ochroptera (Coleoptera: Chrysomelidae) on sesbania

With the widespread introduction of the nitrogen-fixing legume sesbania, Sesbania sesban (L.) Merril, in agroforestry systems, the defoliating beetle Mesoplatys ochroptera St?l has become a serious pest of the trees in Africa. To determine within-field and within-plant spatial distribution of M. ochroptera on both seedlings and trees of sesbania, distribution statistics were computed using Iwao's mean crowding regression model. In 1- to 3-mo-old seedlings, the model accounted for 29.8, 32.2, and 61.0% of the variation observed in mean crowding to mean relationships in egg masses, larvae and adults, respectively. The model slopes of the regression were greater than unity for all stages indicating aggregated spatial distribution. Values of the intercept were greater than zero for egg masses, larvae and adults indicating that the basic components of the population are groups of individuals. The highest density (>80%) of mating and feeding adults was found in the upper third of 1- to 2-mo-old seedlings, while most of the egg masses were found in the lower half of seedlings. In trees, >60% of the individuals of all stages were found in the lower third of the foliage canopy, while <10% were found in the upper third. Sampling adults was found to be easier and gave better density estimates of M. ochroptera population than egg masses and larvae. Therefore, sampling plans useful for population studies and decision-making in pest management were developed for adults.     

Effect of a phloxine B-cucurbitacin bait on diabroticite beetles (Coleoptera: Chrysomelidae)

Cucurbitacin E glycoside, extracted from a bitter mutant of Hawkesbury watermelon [Citrulls lanatus (Thunb.) Matsum. & Nakai (Syn. Citrullus vulgaris Schrad)] is the active ingredient of a feeding stimulant for the corn rootworm complex. It is the primary component of a water-soluble bait that can be combined with toxins for adult diabroticite control. Studies were conducted using phloxine B (D&C Red 28), a xanthene dye, as the toxin. This dye was efficacious against Diabrotica undecimpunctata howardi Barber, spotted cucumber beetle, and Acalymma vittatum (F.), striped cucumber beetle, in cucumber plots and could be recovered from cucumber leaves for 8 d after treatment. The average amount of dye recovered per dead spotted cucumber beetle at 8 d after treatment was 0.173 microg. Concentrated and sugar-free fermented forms of the watermelon extract were developed and compared with the fresh juice in field applications on cucumber plants. There was no significant difference in mortality of beetles from phloxine B-bait prepared with fresh, fermented, or concentrated extract, although in laboratory studies, fermented juice had higher feeding stimulant activity.     

Evaluation of cucurbitacin-based gustatory stimulant to facilitate cucumber beetle (Coleoptera: Chrysomelidae) management with foliar insecticides in melons

The bitter plant-derived compounds cucurbitacins are known to stimulate feeding of adult cucumber beetles (Coleoptera: Chrysomelidae). A cucurbitacin-based gustatory stimulant applied as a flowable bait combined with either spinosad or carbaryl was compared with foliar sprays of spinosad and carbaryl for controlling two cucumber beetle species (Diabrotica undecimpunctata undecimpunctata Mannerheim and Acalymma trivittatum Mannerheim) in honeydew melons (Cucumis melo L.). Field studies were conducted on the University of California-Davis plant pathology farm in 2008 and 2009. Beetle densities after applications and fruit damage from beetle feeding were compared among treatments. In addition, beetle survival was compared within field cages placed over the treated foliage infested with beetles. Using all three measures of efficacy, we determined that the addition of cucurbitacin bait had no effect on the level of cucumber beetle control with carbaryl in either 2008 or 2009. In both years, spinosad did not significantly reduce cucumber beetle densities in either field cages or field plots and did not reduce fruit damage relative to the untreated control. The addition of the bait to spinosad did not improve its efficacy. A laboratory bioassay of the spinosad formulation used in the field showed it had significant lethal effects on adults of both cucumber beetle species. Results indicated that the bait formulation used did not improve cucumber beetle control but may benefit from the addition of floral attractants or using a different type of cucurbitacin.     

Efficiency of flight interception traps for adult Colorado potato beetles (Coleoptera: Chrysomelidae)

Interception traps have been used to monitor and sample Colorado potato beetle, Leptinotarsa decemlineata (Say), populations in flight, but the efficiency of these traps was unknown. The efficiency of interception window traps is limited because flying adults avoid the traps and may bounce off the trap without being collected. All trap types tested were avoided in flight chamber tests, including those constructed of transparent Plexiglas or yellow wood boards. A larger screen trap and a harp trap designed to reduce detection were also avoided by the beetles in flight. None of the traps provide a direct estimate of the number of adults randomly flying in the trap area. The highest level of efficiency for window traps was obtained with the yellow trap, which caught 16% of the expected flying population. Harp and screen traps without frames intercepted 60-62% of the expected flying population. The presence of a frame did not increase significantly the avoidance of the screen and harp traps by beetles in flight. Recommendations are made to maintain or increase the efficiency of some traps.     

Resistance potential of colorado potato beetle (Coleoptera: Chrysomelidae) to novaluron

Novaluron (Rimon 10 EC), a novel insect growth regulator, could play an important role in future management programs for Colorado potato beetle, Leptinotarsa decemlineata (Say). Studies were conducted to determine the potential of Colorado potato beetle to develop resistance to novaluron before its widespread use in Colorado potato beetle management. Second instars of an imidacloprid-resistant Colorado potato beetle strain exhibited reduced susceptibility (2.5-fold) to novaluron. The toxicity of novaluron to this strain was synergized by S,S,S-tributyl phosphorotrithioate (DEF) but not by piperonyl butoxide (PBO), suggesting that esterase-based detoxification mechanisms were responsible for novaluron resistance. Bioassays with treated potato foliage found that a single low- or medium-rate novaluron application was highly persistent under field conditions, resulting in up to 85% mortality of second instars 5 wk after treatment. Thus, intense selection pressure for novaluron-resistant Colorado potato beetle may continue long after population densities have been reduced below an economic threshold level. In a national survey, the susceptibility of second instars to a novaluron diagnostic dose was determined for 27 different field populations collected from six Canadian provinces in summer 2003. Despite no previous exposure to novaluron, mortalities at the diagnostic dose ranged from 55 to 100%. Although novaluron has several characteristics that should delay resistance development in insect pests, these results highlight the need for judicious use of the compound in management of Colorado potato beetle.     

Comparison of adult corn rootworm (Coleoptera: Chrysomelidae) sampling methods

Studies were conducted in Kansas corn and soybean fields during 1997 to compare various sampling methods, traps, and trap components for capturing three species of adult corn rootworms: western (Diabrotica virgifera virgifera Leconte), southern (D. undecimpunctata howardi Barber), and northern (D. barberi Smith & Lawrence). Lure constituents affected the species of beetle attracted to the trap. Traps with a lure containing 4-methoxycinnamaldehyde attracted more western corn rootworms, those with a lure containing eugenol were more attractive to northern corn rootworms, and those containing trans-cinnamaldehyde were most attractive to southern corn rootworms. Multigard sticky traps caught more beetles than did Pherocon AM sticky traps. In corn, a newly designed lure trap caught more beetles than did sticky traps on most occasions. Also, lure-baited sticky traps caught more beetles than did nonbaited sticky traps. Varying the color of the lure trap bottom did not affect the number caught. In soybeans, the new lure traps captured more beetles than did the nonbaited Multigard or Pherocon AM sticky traps. Results of this study suggest the new lure trap may provide a more accurate assessment of corn rootworm populations than traditional monitoring techniques and may be more esthetically pleasing to growers and consultants.     

Comparative analysis of microbial diversity in Longitarsus flea beetles (Coleoptera: Chrysomelidae)

Herbivorous beetles comprise a significant fraction of eukaryotic biodiversity and their plant-feeding adaptations make them notorious agricultural pests. Despite more than a century of research on their ecology and evolution, we know little about the diversity and function of their symbiotic microbial communities. Recent culture-independent molecular studies have shown that insects possess diverse gut microbial communities that appear critical for their survival. In this study, we combined culture-independent methods and high-throughput sequencing strategies to perform a comparative analysis of Longitarsus flea-beetles microbial community diversity (MCD). This genus of beetle herbivores contains host plant specialists and generalists that feed on a diverse array of toxic plants. Using a deep-sequencing approach, we characterized the MCD of eleven Longitarsus species across the genus, several of which represented independent shifts to the same host plant families. Database comparisons found that Longitarsus-associated microbes came from two habitat types: insect guts and the soil rhizosphere. Statistical clustering of the Longitarsus microbial communities found little correlation with the beetle phylogeny, and uncovered discrepancies between bacterial communities extracted directly from beetles and those from frass. A Principal Coordinates Analysis also found some correspondence between beetle MCD and host plant family. Collectively, our data suggest that environmental factors play a dominant role in shaping Longitarsus MCD and that the root-feeding beetle larvae of these insects are inoculated by soil rhizosphere microbes. Future studies will investigate MCD of select Longitarsus species across their geographic ranges and explore the connection between the soil rhizosphere and the beetle MCD.     

Rearing the Mexican beetle Zygogramma bicolorata (Chrysomelidae: Coleoptera) on a semi-synthetic diet

A semi-synthetic diet utilising parthenium leaf powder was evaluated for rearing the Mexican beetle Zygogramma bicolorata Pallister under in vitro conditions at 26±1°C, 65% RH. Significant differences were observed in the biological parameters of the beetle viz., the grub survival (54.3%), pupation (47.2%), fecundity (54.6/day) and total developmental period (37.7 days), when reared on the semi-synthetic diet compared to those when reared on natural parthenium leaves (91.5, 85.5, 74.8 and 29.8, respectively). The feasibility of utilising the semi-synthetic diet for mass production of the beetle is discussed.     

Reproductive activity of Coelomera lanio (Coleoptera: Chrysomelidae)

The Cecropia spp. (Cecropiaceae) trees are attacked by several insects; among them, Coelomera lanio (Dalman) (Coleoptera: Chrysomelidae). The reproductive behavior of C. lanio was studied under laboratory conditions (12 hour photoperiod, 24.1 +/- 0.1 degrees C and mean relative humidity 67.7 +/- 0.6%), in Vi?osa, Minas Gerais, Brazil. The insects were reared in cages and in Petri dishes and fed leaves of Cecropia pachystachya Trec. Reproductive activity began 5.8 +/- 0.2 days after adult emergence and mean copulation time was 2.5 +/- 0.1 min. The female began oviposition only after 25.7 +/- 0.7 days. Each female laid a mean of 4.7 +/- 0.4 times (range 1-9). The mean number of eggs per oviposition and female was 129.2 +/- 2.4 and 587.4 +/- 92.1 respectively, and the time between egg-layings averaged 16.3 +/- 0.8 days.