Ovary structure and transovarial transmission of endosymbiotic microorganisms in Marchalina hellenica (Insecta,Hemiptera, Coccomorpha: Marchalinidae)

Szklarzewicz, T., Kalandyk‐Kolodziejczyk, M., Kot, M. and Michalik, A. 2011. Ovary structure and transovarial transmission of endosymbiotic microorganisms in Marchalina hellenica (Insecta, Hemiptera, Coccomorpha: Marchalinidae). —Acta Zoologica (Stockholm) 00 :1–9. The paired ovaries of Marchalina hellenica are composed of about 200 ovarioles of telotrophic type. In each ovariole, a trophic chamber, vitellarium and ovariolar stalk can be distinguished. The tropharia comprise trophocytes and early previtellogenic oocytes (termed arrested oocytes) or trophocytes only. The arrested oocytes are not capable of further development. In the vitellaria, single oocytes develop that are connected to the tropharium by means of broad nutritive cords. The number of germ cells (trophocytes and oocytes) constituting ovarioles is not constant and may range between 25 and 32. Numerous endosymbiotic bacteria occur in the cytoplasm of trophocytes. The endosymbionts are transported via nutritive cords to the developing oocyte. The obtained results are discussed in a phylogenetic context.     

Differences in bacterial diversity of host-associated populations of Phylloxera notabilis Pergande (Hemiptera: Phylloxeridae) in pecan and water hickory

Host‐associated differentiation (HAD) is the presence of genetically divergent, host‐associated populations. It has been suggested that microbial symbionts of insect herbivores may play a role in HAD by allowing their insect hosts to use different plant species. The objective of this study was to document if host‐associated populations of Phylloxera notabilis Pergande (Hemiptera: Phylloxeridae) in pecan and water hickory corresponded with differences in the composition of their associated bacteria. To test this hypothesis, we characterized the symbionts present in P. notabilis associated with these two tree species through metagenomic analyses using 454 sequencing. Differences in bacterial diversity were found between P. notabilis populations associated with pecan and water hickory. The bacteria, Pantoea agglomerans and Serratia marcescens, were absent in the P. notabilis water hickory population, whereas both species accounted for more than 69.72% of bacterial abundance in the pecan population.     

Structure of ovaries in ensign scale insects,the most primitive representatives of Coccomorpha (Insecta,Hemiptera)

The ovaries of Orthezia urticae and Newsteadia floccosa are paired and composed of numerous short ovarioles. Each ovariole consists of an anterior trophic chamber and a posterior vitellarium that contains one developing oocyte. The trophic chamber contains large nurse cells (trophocytes) and arrested oocytes. The total number of germ cells per ovariole (i.e., cluster) is variable, but it is always higher than 32 and less than 64. This suggests that five successive mitotic cycles of a cystoblast plus additional divisions of individual cells are responsible for the generation of the cluster. Cells of the trophic chamber maintain contact with the oocyte via a relatively broad nutritive cord. The trophic chamber and oocyte are surrounded by somatic cells that constitute the inner epithelial sheath around the former and the follicular epithelium around the latter. Anagenesis of hemipteran ovarioles is discussed in relation to the findings presented. © 1995 Wiley-Liss, Inc.     

毛蚜属Chaitophorus蚜虫(半翅目:蚜科:毛蚜亚科)与初级内共生菌Buchnera的演化关系

【目的】蚜虫体内共生菌种类丰富,二者关系十分密切。几乎所有蚜虫都具有一类专性的初级内共生菌Buchnera aphidicola,二者的专性共生关系使蚜虫-Buchnera成为研究共生关系演化的理想模型。本研究对蚜虫-Buchnera在低级阶元水平上的"平行演化假说"进行了验证。【方法】选取在杨属Populus或柳属Salix植物上营同寄主全周期生活的毛蚜属Chaitophorus蚜虫作为研究对象,基于不同来源的分子标记(蚜虫线粒体基因、核基因和内共生菌基因),运用最大似然法和贝叶斯法重建蚜虫和Buchnera的系统树,并利用Tree Map、Jane和Para Fit检验二者是否具有协同系统发生关系。【结果】Tree Map和Jane分析检测到毛蚜属蚜虫与Buchnera具有显著的共成种信号,Para Fit分析结果表明二者的总体关联极为显著。【结论】毛蚜属蚜虫与其初级内共生菌Buchnera在种级及以下水平上符合"平行演化假说",并且二者的演化关系不会受到寄主植物差异的影响。     

Ultrastructural study of oogenesis in Phoronopsis harmeri (Phoronida)

Temereva, E.N., Malakhov, V.V. and Yushin, V.V. 2011. Ultrastructural study of oogenesis in Phoronopsis harmeri (Phoronida). —Acta Zoologica (Stockholm) 92 : 241–250. The successive stages of oogenesis in Phoronopsis harmeri were examined by electron microscopy methods. During the oogenesis, each oocyte is encircled by vasoperitoneal (coelomic) cells forming a follicle. The previtellogenic oocytes are small cells which accumulate ribosomes for future synthesis; their cytoplasm contains characteristic clusters of mitochondria and osmiophilic particles resembling a germ plasm of other metazoans. The cytoplasm of the vitellogenic oocytes includes numerous mitochondria, cisternae of the rough endoplasmic reticulum, Golgi bodies and annulate lamellae. The synthesis of three types of inclusions was observed: strongly osmiophilic granules (lipid droplets) as a prevalent component, distinctly larger granules surrounded by membrane (proteinaceous yolk) and numerous large vesicles with pale flocculent content. No inclusions which could be unequivocally interpreted as the cortical granules were detected. The surface of the vitellogenic oocytes is covered by microvilli which increase in number and length during development. The oogenesis in Phoronida may be interpreted as follicular because of close association of oocytes with the vasoperitoneal tissue. However, well‐developed synthetic apparatus together with a strongly developed microvillous surface and absence of endocytosis indicate a clear case of autosynthetic vitellogenesis. Thus, in phoronids, there is a combination of simply developed follicle and autosynthesis that, apparently, is plesiomorphic character.     

Organization of the mitochondrial genomes of whiteflies,aphids, and psyllids (Hemiptera,Sternorrhyncha)

Background  

With some exceptions, mitochondria within the class Insecta have the same gene content, and generally, a similar gene order allowing the proposal of an ancestral gene order. The principal exceptions are several orders within the Hemipteroid assemblage including the order Thysanoptera, a sister group of the order Hemiptera. Within the Hemiptera, there are available a number of completely sequenced mitochondrial genomes that have a gene order similar to that of the proposed ancestor. None, however, are available from the suborder Sternorryncha that includes whiteflies, psyllids and aphids.     

Ovarian Structure in Milnesium tardigradum (Tardigrada, Milnesiidae) during Early Vitellogenesis

The ultrastructure of the ovary of Milnesium tardigradum during early vitellogenesis is described. Within the ovary, there were large multinuclear cells surrounded by many mononuclear oocytes. Observation of serial sections revealed four multinuclear cells that were connected to each other by cytoplasmic bridges. Each peripheral oocyte was connected to the multinuclear cell. An enormous ER-like structure was conspicuous in the centre of the multinuclear cell. The presence of large numbers of lipid droplets and yolk granules in both multinuclear cells and many mononuclear oocytes suggested a role as nurse cells. A small number of these oocytes grow to be eggs. The structural features of the multinuclear nurse cell were compared with other known examples.     

Evidence for host-associated clones of grape phylloxera Daktulosphaira vitifoliae (Hemiptera: Phylloxeridae) in Australia

Grape phylloxera, Daktulosphaira vitifoliae Fitch, is an important pest of grapevines (Vitis vinifera L.) (Vitaceae). Using microsatellite DNA markers it was demonstrated strong associations can exist between D. vitifoliae asexual lineages and vine host type within a vineyard. Also, in excised root bioassays, D. vitifoliae collected from three regions where different genotypic classes predominated showed host-specific differences in life table parameters of reproductive rate and intrinsic rate of increase. Lastly, comparisons of mitochondrial DNA (cytochrome oxidase I) sequences revealed that D. vitifoliae in Australia have paraphyletic origins and fall into two clades partially related to vine host usage. These findings indicate introduction of separate lineages of D. vitifoliae which have close host associations and as such, have important implications for management of this pest in Australia.     

Morphological study of the reproductive system and ovarian growth of the univoltine bamboo borer,Omphisa fuscidentalis (Hampson, 1896) (Lepidoptera: Crambidae)

ABSTRACT

The female reproductive system and protein deposition in the ovaries during development have never been examined in the bamboo borer Omphisa fuscidentalis Hampson. The aim of this study was thus to study the morphology of the female reproductive system of each stage of development. The female reproductive system of the borer consists of a pair of ovary, oviduct and accessory glands. Each ovary is composed of four polytrophic ovarioles that connect to lateral oviducts, fused with a common oviduct. The size of the ovary in diapausing larvae for 9 months was determined. The length and width of the ovaries were the smallest in September larvae (0.343 ± 0.03 and 0.071 ± 0.01 mm, respectively). The ovaries were the largest during ovarian development in May (0.752 ± 0.08 mm long and 0.084 ± 0.01 wide). Additionally, ovarian size was significantly larger in adults than in pupae. The ovarian protein concentration of larvae in May was 0.59 ± 0.06 mg/ml and increased to 16.61 ± 7.5 and 37.42 ± 5.5 mg/ml in pupae (June) and adults (July), respectively. The results showed ovarian development in all life stages of this holometabolous insect, which has a longer life cycle than other lepidopterans.

Abbreviation: TEM: transmission electron microscope     

Macroevolutionary patterns in the Aphidini aphids (Hemiptera: Aphididae): diversification, host association, and biogeographic origins

Background

Due to its biogeographic origins and rapid diversification, understanding the tribe Aphidini is key to understanding aphid evolution. Major questions about aphid evolution include origins of host alternation as well as age and patterns of diversification in relation to host plants. To address these questions, we reconstructed the phylogeny of the Aphidini which contains Aphis, the most diverse genus in the family. We used a combined dataset of one nuclear and four mitochondrial DNA regions. A molecular dating approach, calibrated with fossil records, was used to estimate divergence times of these taxa.

Principal Findings

Most generic divergences in Aphidini occurred in the Middle Tertiary, and species-level divergences occurred between the Middle and Late Tertiary. The ancestral state of host use for Aphidini was equivocal with respect to three states: monoecy on trees, heteroecy, and monoecy on grasses. The ancestral state of Rhopalosiphina likely included both heteroecy and monoecy, whereas that of Aphidina was most likely monoecy. The divergence times of aphid lineages at the generic or subgeneric levels are close to those of their primary hosts. The species-level divergences in aphids are consistent with the diversification of the secondary hosts, as a few examples suggest. The biogeographic origin of Aphidini as a whole was equivocal, but the major lineages within Aphidina likely separated into Nearctic, Western Palearctic, and Eastern Palearctic regions.

Conclusions

Most generic divergences in Aphidini occurred in the Middle Tertiary when primary hosts, mainly in the Rosaceae, were diverging, whereas species-level divergences were contemporaneous with diversification of the secondary hosts such as Poaceae in the Middle to Late Tertiary. Our results suggest that evolution of host alternation within Aphidini may have occurred during the Middle Tertiary (Oligocene) when the secondary hosts emerged.     

Forecasting migration of cereal aphids (Hemiptera: Aphididae) in autumn and spring

The migration of cereal aphids and the time of their arrival on winter cereal crops in autumn and spring are of particular importance for plant disease (e.g. barley yellow dwarf virus infection) and related yield losses. In order to identify days with migration potentials in autumn and spring, suction trap data from 29 and 45 case studies (locations and years), respectively, were set‐off against meteorological parameters, focusing on the early immigration periods in autumn (22 September to 1 November) and spring (1 May to 9 June). The number of cereal aphids caught in a suction trap increased with increasing temperature, global radiation and duration of sunshine and decreased with increasing precipitation, relative humidity and wind speed. According to linear regression analyses, the temperature, global radiation and wind speed were most frequently and significantly associated with migration, suggesting that they have a major impact on flight activity. For subsequent model development, suction trap catches from different case studies were pooled and binarily classified as days with or without migration as defined by a certain number of migrating cereal aphids. Linear discriminant analyses of several predictor variables (assessed during light hours of a given day) were then performed based on the binary response variables. Three models were used to predict days with suction trap catches ≥1, ≥4 or ≥10 migrating cereal aphids in autumn. Due to the predominance of Rhopalosiphum padi individuals (99.3% of total cereal aphid catch), no distinction between species (R. padi and Sitobion avenae) was made in autumn. As the suction trap catches were lower and species dominance changed in spring, three further models were developed for analysis of all cereal aphid species, R. padi only, and Metopolophium dirhodum and S. avenae combined in spring. The empirical, cross‐classification and receiver operating characteristic analyses performed for model validation showed different levels of prediction accuracy. Additional datasets selected at random before model construction and parameterization showed that predictions by the six migration models were 33–81% correct. The models are useful for determining when to start field evaluations. Furthermore, they provide information on the size of the migrating aphid population and, thus, on the importance of immigration for early aphid population development in cereal crops in a given season.     

Tibial combs in the Cydnidae (Hemiptera: Heteroptera) and their functional, taxonomic and phylogenetic significance

Tibial combs in representatives of the family Cydnidae are described in detail for the first time. The structure was studied in 98 species of 58 genera representing all the subfamilies, among them 16 species were investigated using scanning electron microscopic (SEM) techniques. In addition, Parastrachia japonensis (Scott, 1880) of the family Parastrachiidae, and two species of Dismegistus Amyot and Serville, 1843 (a genus of uncertain systematic position within Pentatomoidea) have also been studied. Morphological terminology is proposed for all the structures connected with tibial combs and the term 'the tibial comb complex' is suggested; its functional, taxonomic and phylogenetic significance is also discussed. The genera of Cydnidae can be classified into two groups depending on differences in the tibial comb complex, when it is present; moreover, its absence in the tribe Scaptocorini (Cephalocteinae) is demonstrated for the first time and is regarded as an autapomorphy of this tribe. Data on the occurrence of tibial combs in other families of true bugs (Hemiptera: Heteroptera) are also briefly reviewed, and an anagenetic trend in their evolution in terrestrial Heteroptera is hypothesized. The presence of tibial combs on all legs is regarded as the most plesiomorphic state.     

Ecological and genetic aspects of grape phylloxera Daktulosphaira vitifoliae (Hemiptera: Phylloxeridae) performance on rootstock hosts

Performance and genetic variability of clonal lineages derived from one Californian and one German population of grape phylloxera, Daktulosphaira vitifoliae Fitch were studied on their natal grape rootstock host and on three novel hosts over four generations in an aseptic dual culture system. The ability of D. vitifoliae to adapt to new hosts was measured by changes in fitness (rm) over four generations. The performance of a given clonal lineage changed over successive generations, depending upon the host plant and the phylloxera group. Analysis of amplified fragment length polymorphism-polymerase chain reaction (AFLP-PCR) banding patterns from 40 individual parthenogenetic D. vitifoliae revealed equal levels of genetic variation both among the four clonal lineages analysed and within the different generations of one lineage. Analysis of molecular variance (AMOVA) showed no significant differences between the D. vitifoliae lineages reared on different host plants, nor was a correlation between host performance and genotype found.     

Species identification of aphids (Insecta: Hemiptera: Aphididae) through DNA barcodes

A 658-bp fragment of mitochondrial DNA from the 5' region of the mitochondrial cytochrome c oxidase 1 (COI) gene has been adopted as the standard DNA barcode region for animal life. In this study, we test its effectiveness in the discrimination of over 300 species of aphids from more than 130 genera. Most (96%) species were well differentiated, and sequence variation within species was low, averaging just 0.2%. Despite the complex life cycles and parthenogenetic reproduction of aphids, DNA barcodes are an effective tool for identification.     

Evolution of mimicry in the New World Dysdercus (Hemiptera: Pyrrhocoridae)

Evolution of mimicry rings is reconstructed for the New World species of cotton-stainer bugs (Dysdercus, Hemiptera: Pyrrhocoridae). Using a cladistic analysis of 43 species (including D. silaceus Doesburg, 1968 stat. nov . and D. modestus Doesburg, 1968 stat. nov .), evolution of colour patterns and biogeographical history of the clade are hypothesized. Three probable mimicry rings are obtained from the analysis, together with data about their possible model–mimic or comimic–comimic structures: (a) ‘Yellow’ species in north-western South America and in the Andes (several possible models, D. rusticus, D. mimus part., D. imitator part., and D. collaris part. as mimics); (b) ‘Median Yellow–Black’ species with a median-forewing black pattern in north-western South America and Central America (several possible models, D. basialbus as a mimic); (c) ‘Median Red–Black’ species with a median-forewing black pattern in the Caribbean islands (D. andreae, D. discolor, D. neglectus–D. sanguinarius clade, and D. jamaicensis–D. fervidus–D. ocreatus clade as comimics). A few additional, less supported rings are also identified. Two highly polymorphic species (D. obscuratus, D. mimus) seem to enter different mimicry rings in different regions.     

Predicting potential distribution of chestnut phylloxerid (Hemiptera: Phylloxeridae) based on GARP and Maxent ecological niche models

The chestnut phylloxerid, Moritziella castaneivora, has been recently recorded as a forest pest in China. It heavily damaged chestnut trees and has caused serious economic losses in some main chestnut production areas. In order to effectively monitor and manage this pest, it is necessary to investigate its potential geographical distribution worldwide. In this study, we used two ecological niche models, Genetic Algorithm for Rule‐set Production (GARP) and Maximum Entropy (Maxent), along with the geographical distribution of the host plants, Japanese chestnut (Castanea crenata) and Chinese chestnut (Castanea mollissima), to predict the potential geographical distribution of M. castaneivora. The results suggested that the suitable distribution areas based on GARP were general consistent with those based on Maxent, but GARP predicted distribution areas that extended more in size than did Maxent. The results also indicated that the suitable areas for chestnut phylloxerid infestations were mainly restricted to Northeast China (northern Liaoning), East China (southern Shandong, northern Jiangsu and western Anhui), North China (southern Hebei, Beijing and Tianjin), Central China (eastern Hubei and southern Henan), Japan (Kinki, Shikoku and Tohoku) and most parts of the Korean Peninsula. In addition, some provinces of central and western China were predicted to have low suitability or unsuitable areas (e.g. Xinjiang, Qinghai and Tibet). A jackknife test in Maxent showed that the average precipitation in July was the most important environmental variable affecting the distribution of this pest species. Consequently, the study suggests several reasonable regulations and management strategies for avoiding the introduction or invasion of this high‐risk chestnut pest to these potentially suitable areas.     

Oxydemeton-methyl resistance, mechanisms, and associated fitness cost in green peach aphids (Hemiptera: Aphididae)

Susceptibility to oxydemeton-methyl and imidacloprid, and the inhibitory effects of oxydemeton-methyl and some organophosphate compounds on acetylcholinesterase (AChE) and carboxylesterase activity were studied in two populations (Karaj and Rasht) of green peach aphids, Myzus persicae (Sulzer). Results show that the Karaj population was resistant to oxydemeton-methyl but susceptible to imidacloprid. The esterase activity of the resistant and susceptible populations suggests that one of the resistance mechanisms to oxydemeton-methyl was esterase-based. The inhibition assay shows that the AChE of the Karaj population is less sensitive to oxydemeton-methyl and paraoxon derivatives. Regarding the paraoxon derivatives, the smaller paraoxon side chain is more potent against the modified AChE than against the AChE from the susceptible strain. Fertility life table parameters of green peach aphid populations resistant and susceptible to oxydemeton-methyl also were studied under laboratory conditions. The standard errors of the population growth parameters were calculated using the Jackknife method. Results showed that susceptible strain exhibits a significantly higher r(m) than the resistant strain, probably because the resistant strain had a higher generation time than the susceptible strain. These results suggested that the resistant Karaj strain may be less fit than the susceptible strain.     

Virulence of Hypocreales fungi to pecan aphids (Hemiptera: Aphididae) in the laboratory

There is need for efficacious biocontrol agents for aphids in commercial orchards. As a preliminary step to this end we determined the virulence of several Hypocreales fungi to pecan aphids. In the first experiment we tested the virulence of Isaria fumosorosea (ARSEF 3581) blastospores to three pecan aphids Monellia caryella, Melanocallis caryaefoliae, and Monelliopsis pecanis under laboratory conditions. Rates of 1 × 10⁷ or 1 × 10⁸ spores per ml were applied in 2 ml via a spray tower to 90 mm Petri dishes containing 10 aphids each. Mortality and mycosis were determined after 24, 48 and 72 h. Treatment effects were observed by 48 h post-application, and by 72 h the higher application rate caused >90% mortality and mycosis in M. caryella and M. caryaefoliae, whereas <70% was observed in M. pecanis.We conducted two subsequent experiments (Experiments 2 and 3), using the same methodology, to compare the virulence of several Hypocreales species and strains against the aphid of primary economic concern to most pecan growers, M. caryaefoliae. In Experiment 2, we compared blastospores and conidia of two I. fumosorosea strains (ARSEF 3581 and ATCC 20874 [= strain 97]). The blastospores of ARSEF 3581 and conidia of ATCC 20874 showed higher virulence than other treatments and thus were included in Experiment 3, which also compared the virulence of conidia of Beauveria bassiana (GHA strain) and Metarhizium anisopliae (F52 strain). Results in Experiment 3 indicated the highest virulence in I. fumosorosea 3581 blastospores and M. anisopliae (F52) followed by I. fumosorosea (20874) conidia. The detection of pathogenicity to pecan aphids establishes the potential for commercial usage and additional study. Results reported here will narrow treatments to test in future greenhouse and field trials.     

Barcoding aphids (Hemiptera: Aphididae) of the Korean Peninsula: updating the global data set

DNA barcode (mitochondrial COI) sequences are provided for species identification of aphids from the Korean Peninsula. Most (98%) of the 154 species had distinct COI sequences (average 0.05% intraspecific pairwise divergence) relative to the degree of sequence divergence among species (average value 5.84%). For species in common with other regions, barcodes for Korean samples fell near or within known levels of variation. Based on these results, we conclude that DNA barcodes can provide an effective tool for identifying aphid species in such applications as pest management, monitoring and plant quarantine.     

Development of setal patterns in embryonic and adult aphids of the Hormaphidinae （Hemiptera： Aphididae）

The elementary traits and diversity of embryonic setal patterns of 31 species of 22 genera, in three tribes of Hormaphidinae are studied. The embryonic setal patterns to every species are provided. Based on the parsimony principle and the developmental principle, these patterns are analyzed in each tribe following the terminology of Richards （1965）. It is suggested that the setal patterns of abdominal tergites are the most important characters in embryonic and adult setal patterns. In conclusion, neoteny is a phylogenetic trend in the Hormaphidinae. Nipponaphidini and Hormaphidini have more typical neoteny in each genus and more diversity in different types of adult viviparous females than Cerataphidini, by showing more frequent and various protopatterns or mesopatterns.