Identification of the molting hormone of the sweet potato (Bemisia tabaci) and greenhouse (Trialeurodes vaporariorum) whitefly

In order to identify the whitefly molting hormone, whole body extracts of mature 4th instar and newly formed pharate adult Bemisia tabaci (Biotype B) and Trialeurodes vaporariorum were prepared and subjected to reverse phase high performance liquid chromatography (RPHPLC). Ecdysteroid content of fractions was determined by enzymeimmunoassay (EIA). The only detectable ecdysteroids that were present in significant amounts in whitefly extracts were ecdysone and 20-hydroxyecdysone. The concentrations of 20-hydroxyecdysone in B. tabaci and T. vaporariorum extracts, respectively, were 40 and 15 times greater than the concentrations of ecdysone. The identity of the two ecdysteroids was confirmed by normal phase high performance liquid chromatography (NPHPLC). When ecdysteroid content of RPHPLC fractions was assayed by radioimmunoassay (RIA), small amounts of polar ecdysteroids were also detected indicating that these ecdysteroids have a very low affinity for the antiserum used in the EIA. Ecdysteroid at 10.4 mM administered by feeding stimulated 2nd instar whitefly nymphs to molt. Based on our results, it appears that 20-hydroxyecdysone is the whitefly molting hormone.     

Pathogenicity of Aschersonia spp. against whiteflies Bemisia argentifolii and Trialeurodes vaporariorum

Entomopathogenic fungi of the genus Aschersonia are specific for whitefly and scale insects. They can be used as biological control agents against silverleaf whitefly, Bemisia argentifolii and greenhouse whitefly, Trialeurodes vaporariorum. Forty-four isolates of Aschersonia spp. were tested for their ability to sporulate and germinate on semi-artificial media and to infect insect hosts. Seven isolates sporulated poorly (less than 1x10(7) conidia/dry weight) and 10 were not able to infect either of the whitefly species. Several isolates were able to produce capilliconidia. Infection level was not correlated with germination on water agar. After a selection based on spore production and infection, virulence of 31 isolates was evaluated on third instar nymphs of both whitefly species on poinsettia (Euphorbia pulcherrima). Whitefly infection levels varied between 2 and 70%, and infection percentages of B. argentifolii correlated with that of T. vaporariorum. However, mortality was higher for T. vaporariorum than for B. argentifolii, as a result of a higher 'mortality due to unknown causes.' Several isolates, among which unidentified species of Aschersonia originating from Thailand and Malaysia, A. aleyrodis from Colombia, and A. placenta from India showed high spore production on semi-artificial medium and high infection levels of both whitefly species.     

Selection of Beauveria bassiana isolates for control of the whiteflies Bemisia tabaci and Trialeurodes vaporariorum on the basis of their virulence, thermal requirements, and toxicogenic activity

As part of a 3-fold approach to select potential mycoinsecticides for whitefly control, we evaluated infectivity, thermal requirements, and toxicogenic activity of the entomopathogenic fungus Beauveria bassiana (Ascomycota: Clavicipitaceae) under laboratory conditions. Twenty-five native B. bassiana isolates and a commercially available mycoinsecticide (based on B. bassiana) were evaluated for virulence to fourth instar nymphs of sweetpotato whitefly, Bemisia tabaci, and greenhouse whitefly, Trialeurodes vaporariorum, at a concentration of 1 × 10⁷ conidia/ml. All isolates were pathogenic for both whitefly species, whereas mortality rates varied from 3 to 85%. A second series of bioassays was conducted on 10 selected isolates using four 10-fold concentrations ranging from 1 × 10⁵ to 1 × 10⁸ conidia/ml. Median lethal concentrations (LC₅₀) of the four most virulent isolates varied from 1.1 × 10⁵ to 6.2 × 10⁶ conidia/ml and average survival time (AST) of treated nymphs from 5.9 to 7.4 days. T. vaporariorum were significantly more susceptible to all B. bassiana isolates than B. tabaci. The thermal biology of the eight most virulent isolates to both whitefly species was investigated at six temperatures (10–35 °C). The colony radial growth rate was estimated from the slope of the linear regression of colony radius on time and data were then fitted to a modified generalized β function that accounted for 90.5–99.3% of the data variance. Optimum temperatures for extension rate ranged from 23.1 to 27.1 °C, whereas maximum temperatures for fungal growth varied from 31.8 to 36.6 °C. On the basis of their virulence and thermal requirements, three isolates showed promise as candidates for whitefly management in Mediterranean greenhouses. Whilst in vitro production of macromolecular compounds toxic to Galleria mellonella larvae was not a requisite for virulence, ASTs of larvae injected with Sephadex G-25 fractions from candidate isolates ranged from 1.4 to 3.7 days compared with 5–6 days for non-toxic G-25 fractions. In addition, proteinase K treatment significantly reduced their toxic activity suggesting that they were proteins and revealing the potential of these isolates to be further improved through biotechnology to kill the pest more quickly.     

Endosymbiotic bacteria as a source of carotenoids in whiteflies

Although carotenoids serve important biological functions, animals are generally unable to synthesize these pigments and instead obtain them from food. However, many animals, such as sap-feeding insects, may have limited access to carotenoids in their diet, and it was recently shown that aphids have acquired the ability to produce carotenoids by lateral transfer of fungal genes. Whiteflies also contain carotenoids but show no evidence of the fungus-derived genes found in aphids. Because many sap-feeding insects harbour intracellular bacteria, it has long been hypothesized that these endosymbionts could serve as an alternative source of carotenoid biosynthesis. We sequenced the genome of the obligate bacterial endosymbiont Portiera from the whitefly Bemisia tabaci. The genome exhibits typical signatures of obligate endosymbionts in sap-feeding insects, including extensive size reduction (358.2 kb) and enrichment for genes involved in essential amino acid biosynthesis. Unlike other sequenced insect endosymbionts, however, Portiera has bacterial homologues of the fungal carotenoid biosynthesis genes in aphids. Therefore, related lineages of sap-feeding insects appear to have convergently acquired the same functional trait by distinct evolutionary mechanisms—bacterial endosymbiosis versus fungal lateral gene transfer.     

The status ofBemisia tabaci (Hom.: Aleyrodidae),Trialeurodes vaporariorum (Hom.: Aleyrodidae) and their natural enemies in Crete

The non «B» biotype ofBemisia tabaci (Gennadius) is recorded for the first time in Crete in 1992, in the north east and south east of the island.Trialeurodes vaporariorum (Westwood) is the predominant whitefly on plants in the north and west of the island. Three surveys of Crete were made in 1992 and 1993 for natural enemies ofB. tabaci andT. vaporariorum and resulted in the collection of 4 species ofEncarsia, (plus a number of species that are unidentifiable at this time), anEretmocerus sp. (unidentifiable at this time) and a fungal pathogen,Paecilomyces farinosus (Dickson Ex Fries) Brown &; Smith.Encarsia adrianae was identified fromT. vaporariorum; which constitutes its most westerly distribution point and a new host record respectively.B. tabaci andT. vaporariorum were found on horticultural crops, ornamentals and weeds. Populations of both whitefly species were severely depleted on field hosts throughout the island during the winter of 1992/93. Climatic constraints, competition withT. vaporariorum in otherwise suitable niches, effective natural enemies and an observed low level of polyphagy may explain the present limited distribution of the non «B» biotype ofB. tabaci in Crete.     

Difference in feeding behaviors of two invasive whiteflies on host plants with different suitability: implication for competitive displacement

In China, Bemisia tabaci Q (commonly known as biotype Q) has rapidly displaced B (commonly known as biotype B) in the past 6 years. The mechanisms underlying such phenomenon have been studied extensively in recent years; however, we have not come to a definitive conclusion yet. In the present study, the differences in host suitability between B and Q whitefly adults to five host plants (cabbage, cotton, cucumber, poinsettia, and tomato) were evaluated based on their respective feeding behaviors using a direct-current electrical penetration graph (DC-EPG) system. Pair-wise comparisons of B. tabaci B and Q feeding on each of the five host plants clearly indicate that Q feeds better than B on tomato, cotton and poinsettia, while B feeds better than Q on cabbage and cucumber. The EPG parameters related to both phloem and non-phloem phases confirm that cabbage and cucumber are best suited to B, while tomato, cotton, and poinsettia are best suited to Q. Our present results support the contention that host suitability and adult feeding behavior contribute to the competitive displacement of biotype B by biotype Q. The discrepancy between field (previous studies) and laboratory results (this study), however, suggests that 1) whitefly displacement is apparently contributed by multiple factors; and 2) factor(s) other than the host plant suitability may play a vital role in dictating the whitefly biotypes in the field.     

Timing and ecdysteroid regulation of the molt in last instar greenhouse whiteflies (Trialeurodes vaporariorum)

A system of markers has been devised to track the development of 3rd and 4th instar/pharate adult greenhouse whiteflies. Instars were identified based on measurements of body width and body length. Depending upon the host plant, the product of the two measurements was exceptionally useful in distinguishing between instars. Body depth was used to divide the 3rd instar into eight stages and body depth and color and appearance of the developing adult eye were used to divide the 4th instar/pharate adult into nine stages. Under conditions of L:D 16:8 and a temperature of 26±2°C, the body depth of 3rd instars reared on greenbean increased from 0.025 (stage 1) to 0.2 mm (stage 8) and the instar duration was approximately 3 days. The body depth of 4th instars increased from approximately 0.1±0.02 (Stage 1) to 0.3±0.03 mm (Stage 5) and then remained constant or decreased slightly during adult development. Ecdysteroid titers peaked at approximately 120 fg/μg protein during Stages 3 through 6 of the 4th instar. Based on an external examination of developing 4th instars and the fluctuations in ecdysteroid titer, it appears that adult development is initiated in Stage 4 or 5 4th instars. Results from histological studies support this view. In Stage 4 nymphs, a subtle change was observed in the corneagenous cells of the eye. However, most Stage 4 4th instars possessed wing development characteristic of earlier, immature stages. In all Stage 5 insects, wing development had been initiated and the corneagenous cells had become quite distinct. In Stage 6 whiteflies, the wing buds were deeply folded and by Stage 7, spines were observed on the new cuticle, indicating that the adult cuticle was well-formed by this stage. Our study is the first to investigate the timing and regulation of the molt, to monitor ecdysteroid titers in precisely staged 4th instar whiteflies and to examine the internal anatomical changes associated with metamorphosis in these tiny homopteran insects.     

Influence of pesticide treatments on the dynamics of whiteflies and associated parasitoids in snap bean fields

To determine the influence of pesticidetreatments on the population dynamics of thewhiteflies Trialeurodes vaporariorum(Westwood) and Bemisia tabaci Gennadius (Homoptera: Aleyrodidae)and their naturally occurring parasitoids, weperformed field experiments on insecticidesprayed and unsprayed fields during a croppingseason of snap bean in Pradera, Valle delCauca, Colombia. Substantially largerpopulations of whitefly nymphs occurred in theunsprayed field than in the sprayed field. Parasitoids were more frequent in unsprayedthan in sprayed fields with Encarsianigricephala Dozier being more prevalent thanAmitus fuscipennis MacGown & Nebeker. Insprayed fields the nymphs parasitized by E. nigricephala exceeded the unparasitizedwhitefly nymphs at the end of the croppingseason. Our results suggest that while undercurrent agricultural practices whiteflies onsnap beans cannot be exclusively controlled bynaturally occurring parasitoids, parasitoidsmay be integrated with chemical control inorder to reduce crop damage.     

Horizontal transmission of the insect symbiont Rickettsia is plant-mediated

Bacteria in the genus Rickettsia, best known as vertebrate pathogens vectored by blood-feeding arthropods, can also be found in phytophagous insects. The presence of closely related bacterial symbionts in evolutionarily distant arthropod hosts presupposes a means of horizontal transmission, but no mechanism for this transmission has been described. Using a combination of experiments with live insects, molecular analyses and microscopy, we found that Rickettsia were transferred from an insect host (the whitefly Bemisia tabaci) to a plant, moved inside the phloem, and could be acquired by other whiteflies. In one experiment, Rickettsia was transferred from the whitefly host to leaves of cotton, basil and black nightshade, where the bacteria were restricted to the phloem cells of the plant. In another experiment, Rickettsia-free adult whiteflies, physically segregated but sharing a cotton leaf with Rickettsia-plus individuals, acquired the Rickettsia at a high rate. Plants can serve as a reservoir for horizontal transmission of Rickettsia, a mechanism which may explain the occurrence of phylogenetically similar symbionts among unrelated phytophagous insect species. This plant-mediated transmission route may also exist in other insect-symbiont systems and, since symbionts may play a critical role in the ecology and evolution of their hosts, serve as an immediate and powerful tool for accelerated evolution.     

Biological control of thrips and whiteflies by a shared predator: Two pests are better than one

We studied the capacity of one species of predator to control two major pests of greenhouse crops, Western flower thrips (Frankliniella occidentalis (Pergande)) and the greenhouse whitefly (Trialeurodes vaporariorum (Westwood)). In such a one-predator–two-prey system, indirect interactions can occur between the two pest species, such as apparent competition and apparent mutualism. Whereas apparent competition is desired because it brings pest levels down, apparent mutualism is not, because it does the opposite. Because apparent competition and apparent mutualism occurs at different time scales, it is important to investigate the effects of a shared natural enemy on biological control on a time scale relevant for crop growth. We evaluated the control efficacy of the predatory mites Amblyseius swirskii (Athias-Henriot) and Euseius ovalis (Evans) in cucumber crops in greenhouse compartments with only thrips, only whiteflies or both herbivorous insects together. Each of the two predators controlled thrips, but A. swirskii reduced thrips densities the most. There was no effect of the presence of whiteflies on thrips densities. Whitefly control by each of the two predators in absence of thrips was not sufficient, yet better with E. ovalis. However, whitefly densities in presence of thrips were reduced dramatically, especially by A. swirskii. The densities of predators were up to 15 times higher in presence of both pests than in the single-pest treatments. Laboratory experiments with A. swirskii suggest that this is due to a higher juvenile survival and developmental rate on a mixed diet. Hence, better control may be achieved not only because of apparent competition, but also through a positive effect of mixed diets on predator population growth. This latter phenomenon deserves more attention in experimental and theoretical work on biological control and apparent competition.     

Avoidance of natural enemies by adult whiteflies, Bemisia argentifolii, and effects on host plant choice

In this study we asked whether, in the context of a trap crop system, differential predation risks among plants influence host choice patterns of adult whiteflies, Bemisia argentifolii. We investigated whether adult whiteflies avoid natural enemies inhabiting poinsettia (a cash crop) and whether this behavior can be used to increase the movement of whiteflies to cucumber (a trap crop). The potential of cucumber as a trap crop was first evaluated and we found that significantly more whiteflies were attracted to cucumber when the whiteflies were released between the two plants. However, the accumulation of whiteflies on cucumber substantially diminished if the insects had first settled on poinsettia. Under such circumstances, we investigated whether movement of adult whiteflies to cucumber could be increased by creating conditions that would cause the whiteflies that had settled on poinsettia to leave the plant. A mechanical disturbance, consisting of shaking the plant, was first used to test this hypothesis. The shaking caused more whiteflies to leave poinsettia and move onto the trap crop, compared to undisturbed whiteflies. We then asked whether the presence of natural enemies on the cash crop could induce whiteflies to leave the cash crop and move onto the trap crop. Three natural enemies were tested: two predators, Amblyseius swirskii and Delphastus catalinae, and a parasitoid Encarsia formosa. The presence of D. catalinae on poinsettia induced significantly more whiteflies to disperse to cucumber compared to poinsettia with no natural enemies, whereas A. swirskii and E. formosa did not result in a significant increase. Predator avoidance behavior by adult whiteflies should be investigated further in the context of trap cropping and other crop-habitat alterations designed to help manage whitefly abundance.     

Establishment of papaya banker plant system for parasitoid, Encarsia sophia (Hymenoptera: Aphilidae) against Bemisia tabaci (Hemiptera: Aleyrodidae) in greenhouse tomato production

The silverleaf whitefly, Bemisia tabaci biotype B (Gennadius) (Hemiptera: Aleyrodidae), is a key pest of tomato (Solanum lycopersicum L.) and other vegetable crops worldwide. To combat this pest, a non-crop banker plant system was evaluated that employs a parasitoid, Encarsia sophia (Girault & Dodd) (Hymenoptera: Aphelinidae) with whitefly, Trialeurodes variabilis (Quaintance) (Hemiptera: Aleyrodidae), as an alternative host for rearing and dispersal of the parasitoid to the target pest. (a) Multi-choice and no-choice greenhouse experiments were conducted to determine host specificity of T. variabilis to papaya (Carica papaya L.) and three vegetable crops including tomato, green bean (Phaseolus vulgaris L.), and cabbage (Brassica oleracea L.). The result showed that papaya was an excellent non-crop banker plant for supporting the non-pest alternative host, T. variabilis, whose adults had a strong specificity to papaya plants for feeding and oviposition in both multi-choice and no-choice tests. (b) The dispersal ability of E. sophia was investigated from papaya banker plants to tomato and green bean plants infested with B. tabaci, as well as to papaya control plants infested with T. variabilis; and (c) the percent parasitism by E. sophia on T. variabilis reared on papaya plants and on B. tabaci infested on tomato plants was also evaluated. These data proved that E. sophia was able to disperse at least 14.5 m away from papaya plants to target tomato, bean or papaya control plants within 48–96 h. Furthermore, E. sophia was a strong parasitoid of both T. variabilis and B. tabaci. There was no significant difference in percent parasitism by E. sophia on T. variabilis (36.2–47.4%) infested on papaya plants or B. tabaci (29–45.9%) on tomato plants. Thus, a novel banker plant system for the potential management of B. tabaci was established using papaya as a non-crop banker plant to support a non-pest alternative host, T. variabilis for maintaining the parasitoid to control B. tabaci. The established banker plant system should provide growers with a new option for long-term control of B. tabaci in greenhouse vegetable production. Ongoing studies on the papaya banker plant system are being performed in commercial greenhouses.     

Identification and characterization of functional aquaporin water channel protein from alimentary tract of whitefly, Bemisia tabaci

Some hemipteran xylem and phloem-feeding insects have evolved specialized alimentary structures or filter chambers that rapidly transport water for excretion or osmoregulation. In the whitefly, Bemisia tabaci, mass movement of water through opposing alimentary tract tissues within the filter chamber is likely facilitated by an aquaporin protein. B. tabaci aquaporin-1 (BtAQP1) possesses characteristic aquaporin topology and conserved pore-forming residues found in water-specific aquaporins. As predicted for an integral transmembrane protein, recombinant BtAQP1 expressed in cultured insect cells localized within the plasma membrane. BtAQP1 is primarily expressed in early instar nymphs and adults, where in adults it is localized in the filter chamber and hindgut. Xenopus oocytes expressing BtAQP1 were water permeable and mercury-sensitive, both characteristics of classical water-specific aquaporins. These data support the hypothesis that BtAQP1 is a water transport protein within the specialized filter chamber of the alimentary tract and functions to translocate water across tissues for maintenance of osmotic pressure and/or excretion of excess dietary fluid.     

Short neuropeptide F (sNPF) is a stage-specific suppressor for juvenile hormone biosynthesis by corpora allata,and a critical factor for the initiation of insect metamorphosis

Molting and metamorphosis are essential events for arthropod development, and juvenile hormone (JH) and its precursors play critical roles for these events. We examined the regulation of JH biosynthesis by the corpora allata (CA) in Bombyx mori, and found that intact brain-corpora cardiaca (CC)–CA complexes produced a smaller amount of JH than that in CC–CA complexes and CA alone throughout the 4th and 5th (last) instar stadium. The smaller amount of synthesis was due to allatostatin-C (AST-C) produced by the brain. The CC synthesized short neuropeptide F (sNPF) that also suppressed the JH synthesis, but only in day 3 4th stadium and after the last larval ecdysis. For the suppression, both peptides prevented the expression of some of the distinct JH biosynthetic enzymes in the mevalonate pathway. Allatotropin (AT) stimulated sNPF expression in the CC of day 1 5th instar stadium, not of day 3 4th; therefore the stage-specific inhibition of JH synthesis by sNPF was partly due to the stimulative action of AT on the sNPF expression besides the stage-specific expression of the sNPF receptors in the CA, the level of which was high in day 2 4th and day 0 5th instar larvae. The cessation of JH biosynthesis in the last instar larvae is a key event to initiate pupal metamorphosis, and both sNPF and AST-C are key factors in shutting down JH synthesis, along with the decline of ecdysone titer and dopamine.     

B型烟粉虱对寄主转换的适应性

将B型烟粉虱分别从嗜性较强的番茄上转移到嗜性相对较弱的国抗22棉花、泗棉3号棉花和辣椒上, 以及从嗜性较弱的辣椒上转移到嗜性相对较强的番茄、国抗22棉花和泗棉3号棉花上, 观察寄主转移后的F₁代、F₂代、F₃代烟粉虱产卵效应和寄主适应度的变化;将F₄代烟粉虱再转移到原寄主, 观察烟粉虱产卵效应和寄主适应度的恢复情况。结果表明, 烟粉虱在不同嗜性寄主上的产卵效应存在明显的差异。在不同嗜性的寄主之间转移, 烟粉虱的寄主适应度变化趋势不同, 从嗜性较强的寄主转移到嗜好性相对较弱的寄主上, 烟粉虱的寄主适应度迅速下降;从嗜性较弱的寄主向嗜性较强的寄主转移后, 烟粉虱的适应度则会迅速提高。烟粉虱对新寄主的适应速度与其对原寄主和新寄主之间的嗜性差异程度有关, 但一般经过1-2个世代后, 产卵效应会逐渐恢复到烟粉虱在该寄主上的正常水平。从过渡寄主转移到原寄主, 烟粉虱的寄主适应度变化符合一般的寄主转移规律, 但嗜性相对较强的过渡寄主可以刺激烟粉虱提高寄主适应性。     

Particulate surface waxes of whiteflies: morphology, composition and waxing behaviour

ABSTRACT The morphology and lipid composition of wax particles and their application to the body surface of adults were examined in the sweetpotato whitefly, Bemisia tabaci (Gennadius), and the glasshouse whitefly, Trialeurodes vaporariorum (Westwood). The wax particles are extruded from plates found on the abdomen of both species. Their form is determined by the templates (modified microtrichia) from which they are extruded. The emerging ribbons are broken off by rakes on the hind tibiae and applied to the rest of the body in a similar manner for both species. Triacylglycerols comprise 65–75% of the lipid in these particles, with hydrocarbons accounting for 3–7% of the total in B. tabaci and only 0.6-1% in T.vaporariorum. Although distinct compositional differences exist between the species, the hydrocarbons in both are fully saturated and contain n -alkanes and branched molecules having eighteen to forty carbon atoms. Possible roles of these wax particles in reducing water flux, protecting against harmful radiation, and providing a barrier to microorganisms are discussed.     

Similarity in flight activity rhythms in coexisting species of Aleyrodidae, Bemisia tabaci and Trialeurodes abutilonea

Experiments were conducted in Arizona, USA to examine the diel patterns of flight activity of two coexisting species of whiteflies, Bemisia tabaci (Gennadius) and Trialeurodes abutilonea (Haldeman) (Aleyrodidae). Aerial populations were monitored during parts of two growing seasons using sticky traps. It was found that on most days when more than 100 whiteflies were captured, the flight activity rhythms of the two species were remarkably similar. During 13 of 14 weeks if aerial populations of one species were used to predict populations of the other, coefficients of determination were highly significant. This is in part a function of the two species having parallel patterns of pupal eclosion and teneral periods of approximately the same duration. The two species both experience 90% eclosion within 1 h of the onset of photophase and both have a teneral period slightly in excess of 4 h at 27°C. As a result populations of the two whiteflies emerge together, become flight ready, and fly at the same time. Arizona populations of these two introduced species appear to have developed certain very similar behavioral and physiological processes under conditions of a rigorous desert climate.

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Zusammenfassung In Arizona (USA) wurden Experimente durchgeführt, um die Muster der täglichen Flugaktivität von zwei co-existierenden Aleurodidenarten, Bemisia tabaci und Trialeurodes abutilonea zu untersuchen. Mit Hilfe von Klebefallen wurden die Adult-Populationen während zweier Wachstumperioden beobachtet. Die Rhythmen der Flugaktivität während der meisten Tage, an denen mehr als 100 Weisse Fliegen gefangen wurden, waren bemerkenswert ähnlich. Die Bestimmungskoeffizienten waren deutlich signifikant in 13 von 14 Wochen, wenn die Populationen der einen Art benutzt wurde, um die Population der anderen Spezies vorherzusagen. Dies rührt zum Teil daher, dass sich beide Arten in ähnlicher Weise entwicklen. Beide schlüpfen zu 90% innerhalb von 3 Stunden nach Beginn der Photophase und beginnen bei 27°C im Mittel nach 4 Stunden und 12 Minuten zu fliegen. Daraus folgt, dass Populationen beider Arten zur selben Zeit schlüpfen, flugbereit werden und zu fliegen beginnen. Es scheint, dass die in Arizona vorhandenen Populationen dieser beiden eingeführten Arten unter dem Einfluss eines rauhen Wüstenklimas einige sehr ähnliche Verhaltensweisen und physiologische Prozesse entwickelt haben.