Population dynamics of the chestnut gall-wasp,Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae)

Summary As a part of serial study on population dynamics of the chestnut gall-wasp,Dryocosmus kuriphilus, analyses of the distribution of eggs, gall-cells and emergent holes were made from the statistical point of view. Many of distributions of the eggs per bud could be described by the truncated Poisson, but some cases showed slight overdispersion than expected by chance. Because of no linear increase of with increasing (expected mean for complete sample), however, the truncated negative binomial seemed to be not so available for whole series. Distributions of the gall-cells and the emergent holes were, on the other hand, well described by the truncated Poisson distribution when the observed frequency was calculated for respective trees. Negative-binomial tendency found in distributions from some stations consisted of a certain number of trees would be resulted from mixture of Poisson populations with different means. Random or slightly concentrated oviposition and random mortality within galls was thus supposed for future study unless the proof favouring other distribution models would appear.     

Population dynamics of the chestnut gall wasp,Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae)

Summary During the course of a 10-year investigation on the population dynamics of Dryocosmus kuriphilus, a rapid decrease in the number of individuals was noticed in the adult stage. To detect the role of predation by spiders in causing this high adult mortality, spiders collected from the survey station were tested for their reaction to Drycosmus antiserum. About 2050% of the spiders collected during the emergence period of D. kuriphilus reacted positively, showing that the species of Thomisidae, Argiopidae, Agelenidae and Salticidae were the important predators. The predation ratio for the total adult wasp population was estimated at 8.1% in 1968, 20.2% in 1969 and much higher in 1970, increasing with decreasing population density of the wasps. However, the predation by spiders could not account for all of the adult mortality of D. kuriphilus.     

Optimal clutch size of the chestnut gall-wasp,Dryocosmus kuriphilus yasumatsu (Hymenoptera: Cynipidae)

Optimal clutch size of the chestnut gall-wasp, Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae), was examined in galls on wild and resistant chestnut trees in 1988 and 1989. The rate of escape success of newly-emerged adults from galls was an average of 60%, irrespective of cell numbers per gall. Dry mass per cell of a gall (as an index of nutritive condition) decreased with increasing cell number per gall, but was proportional to the mean number of mature eggs of new adults per gall. The number of cells per gall that occurred most frequently did not agree with that attained by the maximum survival rate from young larva to adult emergence of the gall-wasp. This discrepancy was examined from the viewpoint of three factors: 1) quality of offspring, 2) defensive response of the host plant causing mortality of the gall-wasp before cell formation, and 3) fitness per gall vs. fitness per egg. It is concluded that the third factor is most likely to be the one best in explaining the discrepancy.     

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.     

Population dynamics of the chestnut gall-wasp,Dryocosmus kuriphilus Yasumatsu (Hymenoptera: Cynipidae) IV. Further analysis of the distribution of eggs and young larvae in buds using the truncated negative binomial series

Distribution pattern of eggs and the first instar larvae of the chestnut gall-wasp, Dryocosmus kuriphilus, per bud of the chestnut tree was re-examined using the truncatedPoisson and the truncated negative binomial series. The results are as follows:

1. About sixty percent of the distribution are approximated by the truncated negative binomial and another thirty percent by the truncatedPoisson . When the distribution has been approximated by the truncated negative binomial, the show scattered values, some of which are near thePoisson , but the mean value of is about 4 both in eggs and in the first instar larvae.
2. The number of buds which have not been infested by gall-wasps is resulted from various factors as follows: (a) Buds formed after the laying activity of gall-wasps has ceased; (b) Old and shrunk buds which were avoided in laying; (c) Buds in which eggs have died by the time of sampling; and (d) Buds escaping ovipositions by chance. Most of the gall-wasp-free buds located in the middle part of the shoot are accounted for the zero-class expected by the negative binomial series.
3. Brass (1958) moment method for estimating parameters of the truncated negative binomial give good precision within the range of from 1 to 10 and of from 1 to 6.
4. It is concluded that the truncated distributions used are useful for the purpose of analysis of the distribution pattern of the chestnut gall-wasp.

     

Identification of Torymus sinensis and T. beneficus (Hymenoptera: Torymidae), introduced and indigenous parasitoids of the chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae), using the ribosomal ITS2 region

The parasitoid wasps Torymus sinensis and T. beneficus (Hymenoptera: Torymidae) are introduced and indigenous natural enemies, respectively, of the chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae), an invasive pest of chestnuts in Japan. T. beneficus has two emergence types in spring, here tentatively designated as the early-spring strain and late-spring strain. It is very difficult to distinguish these two Torymus species accurately according to their morphological and ecological characteristics. Although the sequences of internal transcribed spacer 2 of the nuclear ribosomal DNA (ITS2) of these parasitoids are very similar, we have developed a pair of primers for amplifying a part of ITS2 that distinguishes species and emergence type. By performing high-resolution electrophoresis of PCR products amplified by this specific primer pair, we have succeeded in accurately identifying T. beneficus (early-spring strain) and some T. sinensis parasitoids. However, this technique was discovered to be inapplicable to identify other T. sinensis and T. beneficus (late-spring strain) parasitoids with the same ITS2 genotype. In spite of this problem, the ITS2 marker appears to be more powerful than any other molecular markers so far reported, since it is applicable to detection of T. beneficus, T. sinensis, and their hybrids.     

The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini)

Abstract Oak gallwasps (Hymenoptera; Cynipidae, tribe Cynipini) are cyclically parthenogenetic insects that induce galls on specific plant hosts in the family Fagaceae. Understanding the processes underlying the evolution of specific oak associations requires knowledge of the phylogenetic relationships among oak gallwasp genera. Although three major lineages of oak gallwasps have been identified, the status and relationships of several species‐poor but biologically significant genera remain unresolved. Two such genera are Chilaspis and Dryocosmus, whose western palaearctic species all gall oaks in the section Cerris. Dryocosmus is particularly significant biologically because it includes: (a) the only palaearctic gallwasp to gall chestnuts, Castanea, and (b) nearctic species. The oak section Cerris is wholly absent from the nearctic, and the relationship between palaearctic and nearctic Dryocosmus is significant for patterns of host plant evolution in the tribe as a whole. We examined the relationships between Chilaspis, Dryocosmus and other oak cynipid genera using cladograms from sequence data for two mitochondrial loci (cytochrome c oxidase subunit I and cytochrome b) and two nuclear loci (the 28S ribosomal gene regions D2 and D3–5). Our analyses support the following conclusions: (1) palaearctic Chilaspis and Dryocosmus species form an intermingled monophyletic group. (2) We propose that Chilaspis Mayr, 1881 is a syn.n. of Dryocosmus Giraud, 1859 and propose the name D. mayri as a comb.rev. for the species previously named C. mayri, and D. nitidus and D. israeli as comb.n. of C. nitida and C. israeli, respectively. (3) We reassess the utility of morphological characters previously regarded as diagnostic for these genera. (4) Two species previously known only from a single generation represent two halves of a single species lifecycle. Dryocosmus nervosus is here designated a syn.n. of D. cerriphilus. (5) The nearctic species D. favus lies outside the palaearctic Chilaspis/Dryocosmus clade, and Dryocosmus as currently recognized is not a monophyletic group. (6) Dryocosmus/Chilaspis is closely related to the other oak gallwasp taxa (Aphelonyx, Plagiotrochus, Pseudoneuroterus, Trichagalma, and some Neuroterus species) galling section Cerris oaks. This implies an early branching evolution of this oak association within this group, and supports previous work showing the rarity of oak gallwasp host shifts.     

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.     

Sources of mortality for a cynipid gall-wasp (Dryocosmus dubiosus (Hymenoptera: Cynipidae)): The importance of the Tannin/Fungus interaction

Summary Emergence success was determined for 1300 galls of the cynipid waspDryocosmus dubiosus. Galls were collected throughout a single host tree (a California coast live oakQuercus agrifolia). Each gall was reared individually in small gelatin capsules. For each gall data was recorded on 17 parameters characterizing hyperparasitism, fungal infestation, leaf tannin levels, inter- and intraspecific competition, and spatial position within the tree. Using contingency table analysis and logistic regression, we determined that the most significant factors influencing the success ofD. dubiosus galls are 1) fungal infestation and 2) chalcid hyperparasitism, both having negative effects. Of the factors investigated we found that leaf tannin level had the strongest influence on the degree of fungal infestation. Fungal infestation, in turn, is lowest in regions of high leaf tannins.     

Fungal community associated with adults of the chestnut gall wasp Dryocosmus kuriphilus after emergence from galls: Taxonomy and functional ecology

The diversity of the fungal community associated with adults of Dryocosmus kuriphilus following emergence was examined using HTS analysis. Ascomycota dominated the fungal core-biome community. The functional guilds of the 90 taxa forming the core-biome were assessed, demonstrating three main groups: saprotrophs, plant pathogens and entomopathogens. Twenty-nine OTUs out of 90 were resolved to species level identifying 26 different fungal species. Among these species, many were cosmopolitan or previously recorded in Europe. Ten taxa were previously recorded on chestnut, including some recognized plant pathogens associated with foliage and green tissues such as Epicoccum nigrum, Gnomoniopsis castanea, Colletotrichum acutatum, Stromatoseptoria castaneicola, Ramularia endophylla. Beauveria bassiana; within the core microbiome, Fusarium larvarum represented the most abundant entomopathogenic species. Some of these species are known to impact directly or indirectly the vitality of the insects in the galls. The chestnut blight pathogen, Cryphonectria parasitica, was never found associated with D. kuriphilus. Based on the present study, an active role for D. kuriphilus as a vector of chestnut fungal endophyte/pathogens cannot be demonstrated but neither ruled out.     

The dormant buds of Rhabdopleura compacta (Hemichordata)

Rhabdopleura has an overwintering stage that consists of two layers of cells surrounding a central yolk mass. This cellular part is surrounded by a thick electron dense capsule which is secreted by the bud itself. The capsule is probably impervious and protective to its contents. Blood vessels join the buds to the zooids of the colony. They form the probable route of transfer of yolk from the zooids to the dormant bud. The capsule of the dormant bud has some structural features in common with the black stolon of the adult zooids. The black stolon is probably formed in a manner similar to that which made the fusellar fabric of the periderm of fossil graptolities.     

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.     

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.     

Negative evidence of parthenogenesis induction by Wolbachia in a gallwasp species, Dryocosmus kuriphilus

The alpha‐proteobacteria of the genus Wolbachia is a widespread group of maternally inherited endosymbionts of arthropod and nematode hosts. Wolbachia infection induces a range of host phenotypes, including cytoplasmic incompatibility, male killing, feminization, and induction of thelytokous parthenogenesis. Heterogony (cyclical parthenogenesis) is a remarkable characteristic of oak gallwasps, Cynipini, the largest tribe of the Cynipidae. A few species of Cynipini are exceptional in that they are univoltine and exhibit thelytokous parthenogenesis, probably because they lost the arrhenotokous generation of their heterogonic ancestor species due to Wolbachia infection. In this study, the presence of Wolbachia was detected using polymerase chain reaction primers for the wsp genes in a thelytokous parthenogenetic species [Dryocosmus kuriphilus (Yasumatsu)] (Hymenoptera: Cynipidae: Cynipini). Approximately 29.8 and 87.1% of adults of the Zhuzhou and Fuzhou strains, respectively, were infected with Wolbachia while all females of the remaining four strains collected from other localities in China were Wolbachia free. The length of the wsp fragment of Zhuzhou and Fuzhou strains was found to be 573 and 561 bp, respectively. The nucleotide sequence of the bacterial wsp fragment indicated that the endosymbiotic bacteria of the Zhuzhou and Fuzhou strains are members of supergroup A, but belong to different clades; they probably originated from two independent infection events. In conclusion, thelytokous parthenogenesis of D. kuriphilus is not caused by Wolbachia infection and the deletion of the arrhenotokous generation is thus not associated with such an infection.     

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.     

Molecular diversity and phylogeography of Andricus curtisii (Hymenoptera,Cynipidae) in Turkey

A 433-bp fragment of the mitochondrial cytochrome b gene and the entire nuclear ITS2 region were sequenced in an oak gall wasp species, Andricus curtisii, which represents 21 populations across its range in Turkey. Forty cytochrome b haplotypes and 63 ITS2 alleles were discovered among the 161 individuals studied. Remarkable genetic variation was detected in the species, with an average haplotype diversity (h) of 0.72 and 0.76 and an average nucleotide diversity (π) of 0.06 and 0.01 for the cyt b gene and the ITS region, respectively. Most populations were genetically distinct, possessing unique cytochrome b haplotypes. Maximum likelihood and Bayesian phylogenetic analyses and application of the haplotype network to both datasets revealed that the most basally located haplotypes/alleles were from the eastern part of Turkey. The detection of older sequences in the phylogenetic trees of the easterly located populations implied an eastern origin of the species. The application of a conventional insect mitochondrial DNA clock to the dataset suggested a splitting of ingroup haplotypes from the outgroup lineages predating the Pleistocene epoch and the formation of two major haplogroups that coincided with the oscillation of the early Pleistocene glaciations.     

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.