Cytological investigations of parthenogenesis in gall wasps (Cynipidae,Hymenoptera)

The chromosome number was determined in eleven heterogonous species of gall-wasps on caks and in four species which reproduce by constant parthenogenesis on wild roses. One of the latter group proved to be a natural triploid. Maturation was studied in representative forms from synapsis through diakinesis and the abortive first division to the second division in the laid egg. Chromosome regulation in the developing egg, the production of parthenogones of different sex and the role of the sperm are discussed briefly.     

Differential rates of invasion in three related alien oak gall wasps (Cynipidae: Hymenoptera)

Abstract. Three related species of oak gall wasps, Andricus corruptrix (Schlechtendal), A. kollari (Hartig) and A. lignicola (Hartig) have entered Britain since the introduction of Turkey oak, Quercus cerris L. in 1735. Their lifecycles involve alternating generations between an agamic generation on the native oak species (Q. petraea, Q. robur and their hybrid Q. x. rosacea), and a smaller, sexual generation on the alien Q. cerris. In examining the distributions of these insects and Q. cerris, we hypothesized that: (1) the invasion will spread more rapidly in places where both host trees are equally abundant than through regions where one of the tree species is substantially less common than the other; (2) interspecific competition between these bud‐galling species will lead to a negative correlation between their abundances at a particular site; (3) differential recruitment of natural enemies from the native hymenopteran fauna will slow the rate of spread in a species‐specific manner. A. kollari arrived nearly 200 years ago and is now found throughout the British Isles, wherever Turkey oak is grown. A. lignicola and A. corruptrix have been here for 30 years, after establishing in S.E. England. A. lignicola is in its final rapid stages of range expansion across England, southern Scotland and N.E. Scotland. A. corruptrix is just beginning to spread through Central and S.W. England. It has occupied proportionally fewer sites behind its invasion front than have the other two species, but is no less abundant at these sites. Nevertheless, distance leaps of up to 50 km were identified in A. lignicola in N.E. Scotland, and the possibility of long‐distance transport of infected trees through the horticulture and forestry trades remains. The co‐occurrence of mature individuals of both host Quercus species does appear to have increased their rates of colonization in A. lignicola and A. corruptrix. There is no evidence, however, to suggest that interspecific competition between the three alien gall formers is an important factor in determining their distributions and abundance within their invaded ranges. All three species have recruited parasitoids and inquilines rapidly from the native fauna; attack rates were highly variable, but showed no evidence of density dependence across sites.     

No host‐associated differentiation in the gall wasp Diplolepis rosae (Hymenoptera: Cynipidae) on three dog rose species

Differences in quality and quantity of secondary compounds, as well as in leaf traits of host plants, may influence the host choice of herbivores. Different host preferences could lead to host‐associated differentiation, the first step of sympatric speciation. In the present study, we investigated whether the rose gall wasp Diplolepis rosae L. (Hymenoptera: Cynipidae) shows genetic differentiation related to its host plants (Rosa canina L., Rosa corymbifera Borkh., and Rosa rubiginosa L.). These three host species radiated recently and subsequently expanded their range. Therefore, we expected a diversification within the closely‐associated phytophagous insects. The process of genetic differentiation should be intensified in D. rosae by its close relationship to the host plant, as well as by its parthenogenetic reproduction (infection rate by Wolbachia sp. of almost 100%). However, using 106 polymorphic amplified fragment length polymorphism markers, we found no genetic differentiation among the wasps from different host plants. The population structuring between geographical localities was also low, suggesting considerable gene flow between sites. In part, the low genetic differentiation between sites is explained by the wide distribution of host species and hybrids between host plants. Hybrids with intermediate traits may facilitate the gene flow between wasp populations exploiting different host species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 369–377.     

No Concordant Phylogeographies of the Rose Gall Wasp Diplolepis rosae (Hymenoptera,Cynipidae) and Two Associated Parasitoids across Europe

According to the Host-tracking Hypothesis, species of higher trophic levels with a close relationship to their hosts, such as parasites or parasitoids, are expected to show spatio-temporal phylogeographic patterns similar to those of their host. Alternatively, with ecological sorting, a subset of the local species pools might shift to a related host species, thereby disengaging common phylogeographic patterns. Here, we compare the phylogeographic structures of the cynipid rose gall wasp Diplolepis rosae across Europe and of two of its most common parasitoids, the wasps Orthopelma mediator and Glyphomerus stigma, by analysing the sequences of two gene fragments (COI and ITS 2). The phylogeographic structures of the three species associated with roses were incongruent. D. rosae had the lowest genetic diversity with one major clade, O. mediator showed the classical phylogeographic structure for Europe with one eastern and one western clade, and G. stigma had the highest diversity but no geographical structuring. This discordance of geographical patterns may be explained by 1) the dispersal propensity of adult parasitoids or 2) the parasitoids having the ability to switch to another host, while the primary host becomes rare or is even not available. Furthermore there was no indication that phylogenetic patterns were affected by Wolbachia infections. Our results document that communities of closely interacting species may be the result of idiosyncratic biogeographic histories.     

The effect of hosptlant and parasitoids on the reproductive success of the parthenogenetic gall wasp Diplolepis rosae (Hymenoptera,Cynipidae)

Summary The univoltine cynipid gall wasp Diplolepis rosae reproduces by an obligate homozygosity promoting system known as gamete duplication. The wasp is confined to roses (Rosa spp) on which it induces large, complex and multichambered galls. In southern Sweden, D. rosae was found to parasitize Rosa canina, R. dumalis, R. rubiginosa, R. villosa, R. sherardi and R. rubrifolia, but not R. majalis and R. rugosa. The distribution of galls shows that there are differences in the relation between wasp and hosplant with respect both to species and individual plants. There is a positive correlation between wasp size and gall (clutch) size. Parasitoid pressure was found to be high, causing D. rosae an estimated average larval loss of approximately 75%, mainly due to the attack of the ichneumonid wasp Orthopelma mediator. The very common cynipid inquiline Periclistus brandtii does not seem to have any negative effects. Overall parasitism and probability of no hatched offspring per gall decrease with increasing gall (clutch) size. The probability of loosing all of a given number of offspring decreases with the number of galls produced. It is suggested that D. rosae, in order to escape parasitoids, needs high ability to establish new colonies. Hence the production of many comparatively small galls, which increases the chance of leaving any offspring, rather than the production of few large galls, maximizing the number of offspring, should be favoured by selection.     

Wolbachia-induced thelytoky in the rose gallwasp Diplolepis spinosissimae (Giraud) (Hymenoptera: Cynipidae), and its consequences on the genetic structure of its host

Cynipids are known to use various reproductive modes (arrhenotoky, thelytoky and strict cyclical parthenogenesis), but the mechanism remains unknown. We have studied the reproductive system of a rose gallwasp, Diplolepis spinosissimae, which was found to exhibit two different reproductive systems according to the population. In eight out of the ten populations studied, all along the Atlantic coast, D. spinosissimae is thelytokous. Males are extremely rare, and all females are homozygous at three microsatellite loci. ''Obligate homozygous parthenogenesis'' was found to be strictly associated with the presence of the endosymbiotic bacterium Wolbachia sp. In the two remaining populations, deprived of Wolbachia, D. spinosissimae reproduced by arrhenotoky as indicated by the larger frequency of males and heterozygosity of females. Allelic diversity, although not zero, was highly reduced in the coastal populations, as a consequence of thelytoky and gamete duplication. We hypothesize that mating of uninfected males with infected females during the first generations after an infection event could explain the small but significant amount of polymorphism observed in those populations. The high level of differentiation indicates a low gene flow, even between geographically close coastal populations.     

Invaders on the move: parasitism in the sexual galls of four alien gall wasps in Britain (Hymenoptera: Cynipidae)

Four alien, host-alternating cynipid gall wasps are established in the British Isles: Andricus kollari, A. lignicola, A. quercuscalicis and A. corruptrix. Their current distributions divide Britain into four zones; all four invaders are present in the south and south-east of Britain, whereas only A. kollari is present in the far north of Scotland. The rank order, according to how far north the distributions of the four invading cynipid species reach, is: A. corruptrix→ A. quercuscalicis→ A. lignicola→ A. kollari. The life cycles of all four cynipid species involve a sexual generation in spring on Quercus cerris, and an agamic generation in autumn on Q. robur. Here we studied the parasitoid attack by four pteromalid species on the sexual generations of the invaders. We collected and reared the galls of all four species at eight sites from the south to the north of the country (two sites per zone). The geographical locations of the sites reflect the invasion history and thus the residence time of the alien species in the four zones. At each site we measured the densities of all host galls on Q. cerris and of both obligatory host-tree species. We also took a series of measures, such as host-tree density and mean host-tree size, to further characterize the tree stands. These measures are referred to as local parameters. Host densities varied between sites and between years. In A. kollari, galling rates were highest in the middle of the country (zone 2) in 1994, whereas in 1995 they increased from the south to the north. In A. lignicola, galling rates in both years were lowest at the sites in zone 3 (closest to its distribution boundary). In A. quercuscalicis, galling rates were found to be lowest at the site most to the north-west in both years, again the one furthest away from the area where this species was first recorded. Mortality caused by parasitoid attack differed from less than 10% to as high as 70% and varied between host species, sites and years. In four out of six cases the historical/regional variables (north/south and east/west) correlated significantly with parasitoid attack rates that were characteristically lowest at sites close to the distribution boundaries. Of the local factors, we found parasitoid attack rates correlated negatively in one case with host density, whereas they correlated positively in four cases with the density of alternative hosts of the parasitoids. In one of the models the local density of Q. cerris trees correlated negatively with parasitoid attack on A. quercuscalicis. For all three host species the terms retained in the minimal adequate models obtained for 1994 and 1995 differed, which might indicate that these communities of native parasitoids and invading host have not yet settled in any definite structure.     

New records of Cynipidae (Hymenoptera) from Italy

New records of Cynipid gall wasps and inquilines for the Italian peninsula and Sicily and their new host plants for the Palaearctic Region are listed and commented on. Among them we find: Cerroneuroterus cerrifloralis (Müllner 1901) as new for Italy and new for the Palaearctic region as host on Quercus suber; Andricus multiplicatus Giraud 1859 on Q. suber, as new host for the Palaearctic region; Aylax papaveris (Perris 1839), reported in Italy over a century ago, but later overlooked; Cerroneuroterus minutulus (Giraud 1859), also reported more than a century ago from Sicily, but later overlooked. Among the inquilines are here listed: Synergus variabilis Mayr 1872, emerged from Janetia cerris (Kollar 1850) galls (Diptera Cecidomyiidae), and found for the first time in the Palaearctic Region as host on Q. suber; Saphonecrus haimi (Mayr 1872) and Saphonecrus barbotini Pujade-Villar & Nieves-Aldrey 1986, are new records for Italy.     

Finding of Torymus koreanus (Hymenoptera: Torymidae) attacking Dryocosmus kuriphilus (Hymenoptera: Cynipidae) in Japan

Torymus koreanus (Hymenoptera: Torymidae) was reared from galls of Dryocosmus kuriphilus (Hymenoptera: Cynipidae) for the first time in Japan. We here report morphological features and partial mtDNA sequencing data of T. koreanus. Torymus koreanus has several common characteristics with species of the cyaneus group defined by Zavada (2003 ). According to the key to species groups defined by Graham and Gijswijt (1998 ), it does not belong to any species group because of the entire posterior margin of metasomal tergum five.     

Status of the subfamily names Charipinae and Alloxystinae (Hymenoptera: Cynipidae)

Abstract. The Alloxystinae (= Allotriinae) comprises three genera ( Alloxysta, Phaenoglyphis and Lytoxysta ) of secondary parasites on aphids. Charipinae comprises a single monobasic genus ( Dilyta # Charips = Glyptoxysta ), a primary parasite of a psyllid. The relevant subfamily and generic synonymies are listed. Lectotypes are designated for Dilyta subclavata Förster and Charips microcera Haliday which are newly placed in synonymy.     

Mild winter temperatures reduce survival and potential fecundity of the goldenrod gall fly, Eurosta solidaginis (Diptera: Tephritidae)

We tested the hypothesis that mild winter temperatures are detrimental to the survival and reproductive potential of insects. We measured survival, body size, and potential fecundity of a freeze tolerant insect, the goldenrod gall fly (Eurosta solidaginis), after overwintering in the laboratory for ~3 mo. frozen at -22 degrees C, unfrozen at 0 degrees C, or unfrozen at 12 degrees C. Larvae held at 12 degrees C suffered high mortality (70%) and relatively low potential fecundity as adults (mean+/-SEM=199+/-11 eggs/female), while those held at 0 degrees C had both low mortality (11%) and high potential fecundity (256+/-15 eggs/female). Freezing (-22 degrees C) increased mortality (30% overall) but did not significantly reduce fecundity (245+/-13 eggs/female). Egg length and width were constant regardless of treatment group or female body size. Analysis of covariance indicated that reduced fecundity in the 12 degrees C group was related to reduced larval body weight following treatment. Patterns of larval weight loss in the experimental treatments were generally correlated with previous reports of latitudinal trends in weight loss through the winter. We conclude that mild winter temperatures may be detrimental to some overwintering insects, particularly species that do not feed following winter diapause. Low temperature and even freezing are beneficial, allowing conservation of energy reserves to maintain high survival and potential fecundity.     

Effect of temperature on development, survival, and fecundity of Microplitis manilae (Hymenoptera: Braconidae)

Microplitis manilae Ashmead (Hymenoptera: Braconidae), a larval parasitoid, is a potential biological control agent of both Spodoptera exigua (Hübner) and Spodoptera litura (F.) (Lepidoptera: Noctuidae). Aspects of the climatic requirements for development, including survival, longevity, and fecundity of M. manilae were studied at six constant temperature regimes (17, 20, 23, 26, 29, and 32°C) in the laboratory. The results showed that developmental duration for egg, larva, pupa, and the entire immature stages shortened in response to temperature increasing from 17 to 32°C. Survival rates of different developmental stages were higher at 20-29°C than at other temperatures. Longevity of M. manilae adults shortened with increasing temperature. The maximum fecundity of M. manilae female equaled 261.0 eggs/female at 26°C. Minimum threshold temperature and effective accumulated temperature for completing a generation of M. manilae were 11.04°C and 205.98 degrees-days, respectively. Both intrinsic rate of increase (r) and finite rate of increase (λ) of M. manilae did not differ between 26 and 29°C, but those were significantly higher at 26 and 29°C than at any other temperatures. The highest net reproduction rate (r(0)) was observed at 26°C, with the value of 97.77, but the lowest was 11.79 at 32°C. These results suggest that the parasitoid is well adapted to temperate and subtropical climates, which implies a significant potential for using M. manilae to control S. exigua because most of areas occupied by these two pests belong to temperate and subtropical regions in southeastern Asia.     

Russian honey bee (Hymenoptera: Apidae) colonies: Acarapis woodi (Acari: Tarsonemidae) infestations and overwintering survival

Honey bee, Apis mellifera L. (Hymenoptera: Apidae), colonies infested by parasitic mites are more prone to suffer from a variety of stresses, including cold temperature. We evaluated the overwintering ability of candidate breeder lines of Russian honey bees, most of which are resistant to both Varroa destructor Anderson & Trueman and Acarapis woodi (Rennie), during 1999-2001. Our results indicate that Russian honey bee colonies (headed by original and supersedure queens) can successfully overwinter in the north, even during adverse weather conditions, owing to their frugal use of food stores and their resistance to tracheal mite infestations. In contrast, colonies of Italian honey bees consumed more food, had more mites, and lost more adult bees than Russian honey bees, even during unusually mild winter conditions.     

Biology of Rhoophilus loewi (Hymenoptera: Cynipoidea: Cynipidae), with implications for the evolution of inquilinism in gall wasps

The sole described indigenous afrotropical cynipid, the South African endemic Rhoophilus loewi , was originally described as a gall inducer. As predicted by phylogenetic relationships, we confirm that R. loewi is instead an inquiline wasp. Although all other cynipid inquilines develop in galls induced by cynipid wasps, or very rarely cecidomyiid midges, R. loewi is the only cynipid inquiline of a lepidopteran gall, inducing secondary inquiline cells in galls induced by a cecidosid moth genus Scyrotis on Rhus species (Anacardiaceae). Rhoophilus is a lethal inquiline; its larval cells expand into the hollow interior of the host gall resulting in death of the gall inducer. Rhoophilus loewi is redescribed and the final-instar larva described for the first time. We describe host-plant associations, the morphology and phenology of gall formation, and the suite of parasitoid Hymenoptera associated with these galls. The community centred on Scyrotis galls is compared with those observed in other hosts of inquiline cynipids. Elucidation of the life history of R. loewi has fundamental implications for understanding the evolution of cynipid wasps.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 153–172.     

Possible pathogens of social wasps (Hymenoptera: Vespidae) and their potential as biological control agents

Abstract

Boutin et al. (2006) claimed that American and Eurasian red squirrels use an unknown environmental cue to anticipate the availability of the abundant food of an autumn seed mast, and produce more young than usual in the previous spring and summer. But these small mammals need increased supplies of protein to produce and support young, therefore they must have had access to some other protein‐rich food that was available before the mast was ripe. There are other small mammalian seed‐eaters that increase their reproductive output ahead of the maturation of a seed mast. It seems likely that, in each case, females are able to produce extra young in advance because they eat the amino acid‐rich inflorescences and unripe seeds of the mast and/or larval insects that also increase their numbers in the spring of a mast year by eating the same enriched plant food.     

A phylogenetic analysis of higher-level gall wasp relationships (Hymenoptera: Cynipidae)

We present the most comprehensive analysis of higher-level relationships in gall wasps conducted thus far. The analysis was based on detailed study of the skeletal morphology of adults, resulting in 164 phylogenetically informative characters, complemented with a few biological characters. Thirty-seven cynipid species from thirty-one genera, including four genera of the apparently monophyletic Cynipini and almost all of the genera in the other tribes, were examined. The outgroup included exemplar species from three successively more distant cynipoid families: Figitidae (the sister group of the Cynipidae), Liopteridae and Ibaliidae. There was considerable homoplasy in the data, but many groupings in the shortest tree were nonetheless well supported, as indicated by bootstrap proportions and decay indices. Partitioning of the data suggested that the high level of homoplasy is characteristic of the Cynipidae and not the result of the amount of available phylogenetically conservative characters being exhausted. The analysis supported the monophyly of the Cynipini (oak gall wasps) which, together with the Rhoditini (the rose gall wasps), Eschatocerini and Pediaspidini formed a larger monophyletic group of gall inducers restricted to woody representatives of the eudicot subclass Rosidae. The inquilines (Synergini) were indicated to be monophyletic, whereas the Aylacini, primarily herb gall inducers, appeared as a paraphyletic assemblage of basal cynipid groups. The shortest tree suggests that the Cynipidae can be divided into three major lineages: one including the inquilines, the Aylacini genera associated with Rosaceae, and Liposthenes ; one consisting entirely of Aylacini genera, among them Aulacidea , Isocolus and Neaylax ; and one comprising the woody rosid gallers (the oak and rose gall wasps and allies), the Phanacis-Timaspis complex and the Aylacini genera associated with Papaveraceae.     

温度对金银花尺蠖存活和繁殖的影响

在室内7个恒温(16、19、22、25、28、31和34℃)条件下研究了温度对金银花尺蠖Heterolocha jinyinhuaphaga Chu生长发育、存活、成虫寿命及繁殖等生物学特性的影响。结果表明,温度对其有显著的影响,金银花尺蠖各虫态的发育历期随温度升高而明显缩短,当温度继续升高到34℃,发育历期反而延长,各虫态发育历期同温度之间呈抛物线变化;25℃时金银花尺蠖各虫态的存活率最高,均达到90%以上,在高温和低温下,存活率则明显下降;低温条件下,金银花尺蠖的雌性比率大于雄性的比率,随着温度的升高,雌性比率在不断的下降;成虫寿命与温度成负相关,随着温度的升高而缩短;平均单雌产卵量在25℃时最高,为232.34粒,高温和低温下,则明显下降。