Oviposition behaviour of two tephritid fruit flies,Dacus tryoni and Dacus jarvisi,as influenced by the presence of larvae in the host fruit

Summary When offered a choice, females of the fruit flies Dacus tryoni (Frogg.) and D. jarvisi (Tryon) strongly preferred to lay in fruits without larvae rather than fruits which already contained larvae. Fruits which contained even low densities of larvae, including newly hatched ones, received many fewer eggs than control fruits. This preference was not influenced by the species of larvae present in the fruits nor by the distance to uninfested fruits. Discrimination occurred when fruits with and without larvae were close together (10 cm apart) and also when they were separated by distances of about one metre. Laboratory assays suggested that the flies detect chemical changes in the fruit associated with the decomposition which accompanies larval feeding, but they do not seem to detect the larvae Perse. This behaviour may be significant when these two species utilise the same host since the species which is able to infest fruits first will reduce the availability of hosts for the other species. In contrast to many other Tephritids (e.g. Rhagoletis, Anastrepha and Ceratitis) female Dacus don't discriminate against fruits which contain eggs nor do they deposit a pheromone to deter oviposition by females that subsequently visit the fruit. An hypothesis is proposed to explain the absence of oviposition-deterring pheromones in Dacus, and their presence in many other species of Tephritidae, on the basis of differences in life history and population structure.     

The gene <Emphasis Type="Italic">transformer-2</Emphasis> of <Emphasis Type="Italic">Anastrepha</Emphasis>fruit flies (Diptera,Tephritidae) and its evolution in insects

Background  

In the tephritids Ceratitis, Bactrocera and Anastrepha, the gene transformer provides the memory device for sex determination via its auto-regulation; only in females is functional Tra protein produced. To date, the isolation and characterisation of the gene transformer-2 in the tephritids has only been undertaken in Ceratitis, and it has been shown that its function is required for the female-specific splicing of doublesex and transformer pre-mRNA. It therefore participates in transformer auto-regulatory function. In this work, the characterisation of this gene in eleven tephritid species belonging to the less extensively analysed genus Anastrepha was undertaken in order to throw light on the evolution of transformer-2.     

A high‐throughput detection method for invasive fruit fly (Diptera: Tephritidae) species based on microfluidic dynamic array

Invasive species can be detrimental to a nation's ecology, economy and human health. Rapid and accurate diagnostics are critical to limit the establishment and spread of exotic organisms. The increasing rate of biological invasions relative to the taxonomic expertise available generates a demand for high‐throughput, DNA‐based diagnostics methods for identification. We designed species‐specific qPCR primer and probe combinations for 27 economically important tephritidae species in six genera (Anastrepha, Bactrocera, Carpomya, Ceratitis, Dacus and Rhagoletis) based on 935 COI DNA barcode haplotypes from 181 fruit fly species publically available in BOLD, and then tested the specificity for each primer pair and probe through qPCR of 35 of those species. We then developed a standardization reaction system for detecting the 27 target species based on a microfluidic dynamic array and also applied the method to identify unknown immature samples from port interceptions and field monitoring. This method led to a specific and simultaneous detection for all 27 species in 7.5 h, using only 0.2 μL of reaction system in each reaction chamber. The approach successfully discriminated among species within complexes that had genetic similarities of up to 98.48%, while it also identified all immature samples consistent with the subsequent results of morphological examination of adults which were reared from larvae of cohorts from the same samples. We present an accurate, rapid and high‐throughput innovative approach for detecting fruit flies of quarantine concern. This is a new method which has broad potential to be one of international standards for plant quarantine and invasive species detection.     

X-ray doses to safely release the parasitoid Diachasmimorpha longicaudata (Hymenoptera: Braconidae) reared on Anastrepha fraterculus larvae (Diptera: Tephritidae)

Diachasmimorpha longicaudata is a koinobiont larval parasitoid that is currently used to control fruit flies of the genera Anastrepha, Ceratitis and Bactrocera. In the rearing process, a fraction of the host larvae that are exposed to parasitoids escape from parasitism and develop into viable and fertile flies. This creates the need to eliminate emerging flies before the parasitoids are shipped for release, increasing costs due to additional handling steps. Exposure of fly eggs or larvae to gamma-irradiation before they are parasitised has been used to reproductively sterilise hosts, or even inhibit their emergence. Our aim was to determine whether X-ray radiation applied to Anastrepha fraterculus third instar larvae before they are exposed to parasitoids, inhibits fly emergence in non-parasitised larvae without affecting the performance of the parasitoids that emerge from parasitised larvae. Three X-ray doses: 6250.2 R, 8333.6 R and 10417 R (equivalent to 60, 80 and 100 Gy, respectively) and one γ-ray dose (100 Gy) were tested. Fly emergence decreased with increasing doses of radiation, showing null values for the higher X-ray dose and the dose of 100 Gy. Irradiation showed either no impact or a positive effect on parasitism rate and fecundity. Sex rate was biased towards females in almost every dose. We conclude that the two types of radiation evaluated here were equally effective in suppressing fly emergence with no detrimental effects on the biological quality of the produced parasitoids. X-rays offer an alternative method of irradiation than the conventional radiation source, i.e. γ-rays. These results represent a significant improvement in the development of a biological control programme against A. fraterculus.     

Facultative polar extrusion of yolk masses and of hatching at the posterior egg pole in tephritid fruit flies (Diptera)

The eggs of Anastrepha fracterculus, A. sororcula, A. obliqua, A. serpentina and of Ceratitis capitata (Diptera : Tephritidae) show considerable morphological differences. Differences were also observed in the depth the eggs are inserted into the host fruits. In vivo inspection of the embryos of Anastrepha species showed that they may extrude polar masses of yolk before dorsal closure. In A.fracterculus and A. sororcula, 4 classes of embryos were found : (1) yolk masses at both ends; (2) masses only at the anterior end; (3) just a posterior mass and (4) embryos without any free mass. Only embryos of classes 3 and 4 were found in A. obliqua, while in A. serpentina, all the embryos extruded masses by the posterior end (class 3). There are variations in the frequencies of the classes as well as in the size of the extruded masses. Just before eclosion, the larvae of the Anastrepha species ingest the anterior mass of yolk (when present), turn round inside the egg, suck the posterior extruded mass, and eclose near the posterior-third of the eggs. No masses were present in samples of Ceratitis capitata embryos and the larvae do not present these complex behaviors and eclose near the anterior pole of the egg. The few previous studies on the embryology of other tephritid species did not report similar phenomena. The differences found among the species in relation to the phenomenon of yolk mass extrusion correlate, apparently, to the taxonomic relationship among them, as established before by genetic and morphologic parameters. Additional analyses of other species and genera are required to confirm if yolk mass extrusion is distinctive for the genus Anastrepha.     

The Distributions of Parasitoids (Hymenoptera) of Anastrepha Fruit Flies (Diptera: Tephritidae) along an Altitudinal Gradient in Veracruz, Mexico

In the state of Veracruz, Mexico, fruits from 38 sites at various altitudes were collected monthly over a period of 2 years, and the tephritid fruit flies of the genus Anastrepha and associated parasitoids that emerged from these fruits were identified and counted. Of the 26 species of fruits that contained Anastrepha larvae, 18 species also contained a total of 10 species of Anastrepha parasitoids. These consisted of 4 native and 1 exotic species of opiine braconid larval–pupal parasitoids, 2 native species of eucoilid larval–pupal parasitoids, 1 exotic species of eulophid larval–pupal parasitoid, 1 exotic species of pteromalid pupal parasitoid, and 1 native species of diapriid pupal parasitoid. Overall parasitism (including flies from fruit species that bore no parasitoids) was 6% and was greatest, 16%, at 600–800 m in altitude. The relative contributions of individual parasitoid species to overall parasitism were frequently influenced by both the altitude (and correlated changes in temperature and precipitation) and the species of plant in which the Anastrepha larvae were found. This was particularly the case among the more abundant and widespread Braconidae. To distinguish the role of altitude from that of the distributions of the host plants, these braconids were examined in 4 individual species of fruit that grew over a broad range of altitudes. In guava (Psidium guajava L.) and “jobo” (Spondias mombin L.) the parasitoid Doryctobracon areolatus (Szepligeti) was relatively more common at low altitudes. Its congener, Doryctobracon crawfordi (Viereck), was relatively more abundant at high altitudes in sour orange (Citrus aurantium L.). Utetes anastrephae (Viereck) became relatively more common at higher altitudes in S. mombin, whereas Diachasmimorpha longicaudata (Ashmead) tended to become relatively rare at the highest altitudes in C. aurantium, but increased at high altitudes in P. guajava compared to other braconids. Different altitudinal patterns of abundance in different fruits suggests the importance of both biotic and abiotic factors in parasitoid distributions. We discuss the effect of an expanding agricultural frontier on parasitoid abundance and relate our findings to the design of a fruit fly biological control program that tailors mass releases to parasitoid climate preferences.     

The ovipositor musculature of Carpomya schineri (Loew) (Diptera, Tephritidae)

The structure of the ovipositor sclerites and musculature was studied in the tephritid fly Carpomya schineri (Loew) whose larvae develop in the fruits of Rosa. The structural characters of the ovipositor of this species facilitating laying eggs into such fruits are discussed. The ovipositor musculature and functioning in Carpomya are very similar to those in Ceratitis and Bactrocera; species of all these genera lay eggs in fruits. In the details of the ovipositor musculature, Carpomya is more similar to Ceratitis than to Bactrocera. The genera Carpomya, Ceratitis and Bactrocera are phylogenetically close but Ceratitis and Bactrocera belong to sister tribes within one subfamily, Dacinae. Thus, morpho-functional similarity of the ovipositor structure may not only reflect close relationship but also result from common adaptive transformations.     

Biological attributes of three introduced parasitoids as natural enemies of fruit flies,genus Anastrepha (Diptera: Tephritidae)

The biological attributes of three introduced species of parasitoids which attack the fruit fly Anastrepha ludens were evaluated. Larvae and eggs of A. ludens were exposed to larval parasitoids Diachasmimorpha longicaudata and D. tryoni and the egg parasitoid Fopius arisanus. Parasitoid longevity and fecundity were determined using larvae and eggs of A. ludens. Likewise, the parasitism rates of these parasitoid species in infested host fruits were recorded. The intrinsic rate of increase for F. arisanus was 0.1019 followed by D. tryoni with a rate of 0.1641 and D. longicaudata with the highest rate of 0.2233. Although F. arisanus females had the highest levels of fecundity, only 50% of them remained alive until reproductive age. These results in combination with the longer generation time (in comparison with D. longicaudata and D. tryoni), can be considered as the most important factors explaining F. arisanus reduced rate of increase. However, we note that oviposition activity caused egg mortality which reduced Anastrepha egg hatch by ca. 20%. This result suggests that F. arisanus has a high potential as a natural enemy of A. ludens, in accordance with our research efforts to develop a new F. arisanus strain specialized for development in Anastrepha eggs. The results show that D. tryoni is not a good candidate for biological control of Anastrepha. Meanwhile, D. longicaudata continues to be the most important exotic parasitoid for suppression of Anastrepha fruit fly populations.     

Description of third instar larvae of Ceratitis fasciventris,C. anonae,C. rosa (FAR complex) and C. capitata (Diptera,Tephritidae)

Third instar larvae of members of the Ceratitis FAR complex, including Ceratitis fasciventris (Bezzi), Ceratitis anonae Graham, and Ceratitis rosa Karsch are described and compared with those of Ceratitis capitata (Wiedemann). Diagnostic characters, such as presence vs. absence of a secondary tooth on the mandibles, previously used to separate Ceratitis capitata from Ceratitis rosa, are shown to vary in each species. Significant variation in diagnostic morphological characters among populations of Ceratitis rosa from east and south Africa is documented; however, the differences are not simply congruent with the R1 and R2 designations based on other studies. Quantitative measures of numerous morphological characters are consistently smaller in the larvae of Ceratitis fasciventris and distinguish them from other species of the FAR complex. Larvae of Ceratitis capitata can be distinguished from those of the FAR complex by characters such as absence of accessory plates of the oral ridges, the shape of the anterior spiracle, and the pattern of dorsal spinules. Previous studies indicated that absence of accessory lobes separate the genus Ceratitis from Bactrocera, but this is shown to be incorrect, as accessory lobes are in fact present in several species of Ceratitis.     

我国口岸小条实蝇属截获情况分析

[目的] 小条实蝇属害虫原产非洲,在我国无分布且具有一定的入侵风险。分析该虫在我国口岸的截获情况,能更好地利用已知实蝇截获数据防范实蝇人为传播。[方法] 经动植物检验检疫信息资源共享服务平台查询,统计小条实蝇属的有害生物类别、截获途径及截获地区等。[结果] 全国口岸共截获8种共计524批次的小条实蝇属害虫;地中海实蝇的截获批次数最多,为415批次,为小条实蝇属截获率的79.2%;截获的种类最多的国家来自尼日利亚,5种;截获批次最多的国家为南非,71批次;截获方式最多的为旅客携带物,占总批次的68.3%;在我国口岸中,广东口岸截获批次最多,占总数的57.4%。[结论] 从今后防控的重点种类、加大查验力度、推广新技术在鉴定上的应用、应急预案等几个方面,加强针对该类有害生物的检验检疫措施。     

Sex-specific glands in tephritid fruit flies of the genera Anastrepha,Ceratitis, Dacus and Rhagoletis (Diptera : Tephritidae)

The objectives of this work were to systematically examine males and females of tephritid fruit flies from the genera Anastrepha, Ceratitis, Dacus and Rhagoletis for the presence of possible sex pheromone glands, and to determine whether particular glands may be characteristic of certain genera. Flies from the 4 genera were examined from paraffin embedded histological sections. Pleural epidermal glands and sexually dimorphic, enlarged salivary glands were found in males, but not in females, of 7 species of Anastrepha and in 2 species of Ceratitis. Females of these species do not develop the pleural epidermal glands at all, and although they have salivary glands, the glands do not have the large glandular mass of tissue that is found in males. The pleural glands were similar in all the males, and consisted of a single layer of enlarged, columnar epidermal cells in the pleural region of abdominal segments 3, 4 and 5. The salivary glands of males were divided into 4 types based upon structure.Pleural glands and enlarged salivary glands were searched for, but were not found, in 2 species of Rhagoletis and in 3 species of Dacus. Anal glands, opening externally on the last segment, were confirmed in C. capitata, and were found in C. rosa. Anal glands were not found in any of the other species. A diverticulum from the rectum, the rectal pouch or gland, was confirmed in D. dorsalis, D. cucurbitae and D. tryoni as already published, but this gland was not found in species of other genera. Growth of the pleural glands and salivary glands in males of A. ludens, A. fraterculus and C. capitata was described as a function of age of flies.     

Discovery of novel trimethylalkanes in the internal hydrocarbons of developing pupae of Heliothis virescens and Helicoverpa zea

Novel trimethyl-branched alkanes which eluted with the monomethylalkanes were identified in the internal lipids of Helicoverpa zea but were not present in Heliothis virescens. Their structures were unique in that the first methyl branch occurred on carbon 2 and the 2nd and 3rd methyl branch points were separated by a single methylene. Novel trimethylalkanes identified from their chemical ionization and electron impact mass spectra were 2,18,20-trimethyltetratriacontane, 2,18,20-trimethylhexatriacontane, and 2,24,26-trimethyldotetracontane. Previous reports did not find these trimethylalkanes in the cuticular surface lipids of larvae, pupae or adults of either species. The internal pupal hydrocarbons of H. virescens and H. zea amounted to 123 μg and 304 μg per pupa, respectively. They consisted of n-alkanes (8 and 4%, respectively) and methyl-branched alkanes (88 and 94%, respectively). The n-alkanes ranged in chain length from approximately 21 to 35 carbons and the methyl-branched alkanes from approximately 26 to 55 carbons vs. methyl-branched alkanes from 28 to 37 carbons previously reported for hydrocarbons from the pupal cuticular surface. The major n-alkane was heptacosane (3.3 and 1.2%, respectively, in H. virescens and H. zea). The major methyl-branched alkanes in H. virescens were methylhentriacontane (15%), methyltritriacontane (12%) and dimethyltritriacontane (10%), and in H. zea were methylnonacosane (17%), dimethylnonacosane (9%) and methylhentriacontane (20%). Except for the novel trimethylalkanes, the methylalkane branch points were predominantly on odd-numbered carbons as has been reported for these and other species.     

Size specific demography of three species of Anastrepha fruit flies

Individuals of three Anastrepha species: A. obliqua, A. ludens, and A. serpentina (Diptera: Tephritidae), were sorted according to pupal weight in cohorts of large and small flies. Demographic parameters and reproductive patterns and heterogeneity were determined for each cohort. Large flies of the three species presented greater expectation of life and gross fecundity rates. A. ludens was the species with the longest life span (expectation of life of large adults was 110 days) and the greatest gross fecundity rates (1597 eggs/female for the large flies). While, A. obliqua had the shortest mean age of reproduction (33 days), and the greatest daily egg production (14 eggs/female/day). Net fecundity was similar in these two. A. serpentina had lower fecundity rates.Reproductive information for each size and each species include: age-by-parity relations, fraction of sexually mature life in which females lay eggs, and frequency distribution of individual egg production. Results demonstrate that even under constant laboratory conditions and using standard artificial hosts, there is a great deal of life history variation among these Anastrepha species and among other tephritid fruit flies.     

褐梗天牛幼虫和成虫空间分布的地统计学研究

褐梗天牛(Arhopalus rusticus Linnaeus)是一种对松、杉、柏等林木危害非常严重的蛀干害虫,主要危害针叶树的衰弱木以及火灾后的枯立木,是继松褐天牛之后携带拟松材线虫能力最强的蛀干害虫。为了更好地控制其危害,深入地研究其种群空间格局,运用地统计学方法分析了3块受害程度不同的油松林内褐梗天牛幼虫和成虫的空间分布特性。结果表明:3种受害程度不同的林分内,褐梗天牛的危害具有显著差异,轻度受害林分内有虫株率为30.8%,中度受害林分内有虫株率为44.3%,而重度受害林分内高达78.3%。根据变异函数曲线图分析得知:轻度受害林、中度受害林和重度受害林内褐梗天牛幼虫种群空间分布最优拟合模型分别为高斯模型、高斯模型和指数模型,成虫的种群空间分布最优拟合模型均为线性模型。在3种林分中褐梗天牛幼虫数量具有明显的空间依赖性,轻度受害林、中度受害林和重度受害林内幼虫数量的空间依赖范围分别是19.10、11.97、61.98m,其空间连续性强度分别是0.646、0.784和0.500;成虫的空间依赖范围分别是43.08、43.23、44.17m,其空间连续性强度分别是0.044、0.021和0.171,但其成虫的数量在空间呈随机分布,没有表现出空间依赖性。根据垂直分布图分析得出:褐梗天牛幼虫和成虫在油松上主要集中聚集在某个高,然后随着高度增加密度降低,随着高度接近地面密度也降低。用Kriging插值法生成的空间分布图显示幼虫在空间分布上具有明显的聚集性,其聚集中心主要集中在林地中心,由林地中心株向整个林地扩散,而成虫则表现为随机分布。     

Histone content in relation to amount of heterochromatin and developmental stage in three species of Drosophila

Relative amounts of various histone fractions in Drosophila chromatin were estimated densitometrically on electrophoretic gel separations. Several consistent and highly significant differences were obtained between larval and adult chromatin. The arginine-rich histones showed the most conspicuous changes: higher amounts of H4 in larvae, higher H3 in adults. The level of modification of these histones was clearly higher in larval than in adult chromatin. The modification of the two slower subfractions of H4 involved, in all probability, phosphorylation as well as acetylation. In all types of Drosophila chromatin studied 50% or more of the H2a molecules were phosphorylated—a remarkably high proportion. The species differences observed in relative amounts of histone were consistent in both stages of development. D. melanogaster differed from D. hydei and D. virilis in all histones except H2b, while the latter two species were generally similar. The interspecific variation in histone pattern was generally not correlated to differences in content of heterochromatin. The level of modification of H3 was, however, presumably an exception, as it was significantly lower for both larvae and adults in D. virilis than in the other two species. These differ from D. virilis in containing appreciably lower proportions of heterochromatic chromosome segments.     

Musculature of the ovipositor of Lenitovena trigona (Matsumura) (Diptera, Tephritidae) with reference to the larval saprophagy

The larvae of most Tephritidae are endobiotic in various living plants and are therefore convenient objects for analysis of the associations of this dipteran family with seed plants. The structure of the sclerites and musculature of the ovipositor of Lenitovena trigona (Matsumura) a Far Eastern tephritid of the tribe Acanthonevrini with initially saprobiont larva, was investigated to facilitate understanding of the morphological changes in the adults associated with larval shift from saprophagous to herbivorous habit. Differences between the ovipositor of L. trigona and those of species of the genera Urophora, Ceratitis, and Rhagoletis with endophytic larvae have been found; the ovipositor musculature of these genera has been described by Dean (1935), Nanna (1938), and Berube and Zacharuk (1983).     

Respiratory metabolism of Hydromedion sparsutum and Perimylops antarcticus (Col., Perimylopidae) from South Georgia

Summary Metabolic rates were studied in adults and larvae of the two phytophagous beetles Hydromedion sparsutum and Perimylops antarcticus (Col., Perimylopidae) indigenous to the Sub-Antarctic island of South Georgia. From the regression lines for log₁₀ metabolic rate on temperature no difference in oxygen consumption was found between adults and larvae of Hydromedion or adults of Perimylops from lower sites near sea level and upper sites at about 200 m. Elevated rates of metabolism, however, was found in larvae of Perimylops from upper sites. The slopes of the regression lines of corresponding upper and lower sites adults or larvae were similar in all cases. The metabolic rates were of the same level in adults of both species and in the larvae, but generally higher in adults than in larvae. The activation energies, calculated from Arrhenius plots, varied from 6.7 to 13.6 kcal.mol^–1 in larvae and from 10.7 to 11.3 kcal.mol^–1 in adults. The low values may be interpreted as an expression of cold adapted metabolism. Compared with published data on phytophagous beetles, the metabolic rates of the two species from South Georgia are comparable to two alpine species, but lower than the rates of an Arctic species.     

Susceptibility of Delia radicum to steinernematid nematodes

Isolates of different Steinernema species (S. affine, S. bicornutum, S. feltiae and Steinernema C1) were used in mortality assays with third instar larvae of Delia radicum (L.) (Diptera: Anthomyiidae). The nematode isolates had been obtained by baiting soil regularly grown with cabbage. One isolate (S. feltiae) was the result of a natural infection of a D. radicum puparium. The highest mortality (77%) was obtained with an isolate of S. feltiae (DK1). The isolate DK1 was also used in tests with all larval stages of D. radicum. Mortality around 60% was observed for second and third instar larvae, while first instar larvae showed very low or no susceptibility. Maximum mortality of second and third instar larvae was reached applying only 25 nematodes per larva. Observations of larvae that pupated revealed that some of these puparia contained nematodes. Experiments with hatching puparia showed that a high proportion was infected by nematodes if the flies were prevented from leaving nematode-containing soil. In addition to mortality, the ability of the nematodes to successfully reproduce in the insects was studied. It was found that the species S. feltiae and S. bicornutum reproduced in D. radicum larvae and adults with S. feltiae being the most successful.