Structure and mechanics of the tarsal chain in the hornet, Vespa crabro (Hymenoptera: Vespidae): implications on the attachment mechanism

Two combined mechanisms on the hornet tarsus are adapted to attachment to the substrate: a friction-based (claws and spines) and an adhesion-based one (arolium). There are two ranges of substrate roughness optimal for attachment, either very smooth or very rough. There is an intermediate range of substrate grains of small but non-zero size, where both of these mechanisms fail. The optimal size of substrate grains for hornet grasping was 50-100 microm. Maximal hold to the substrate was achieved when surface irregularities were clamped between the claws of opposite legs. In such a position, the insect could withstand an external force which was almost 25 times larger than its own weight. The tarsal chain is an important part of the entire attachment mechanism. The articulations in the kinematic chain of tibia-tarsus-pretarsus are monocondylar. Three tarsal muscles and one head of the claw retractor muscle originate in the tibia. On pull to the retractor tendon, the tarsus bends in a plane. All elements of the tarsal kinematic chain have one active degree of freedom. The distance between the intertarsomeric articulation point and the tendon of the claw retractor (75-194 microm) corresponds to an efficiency of 1 degrees per 1-3 mircom of pulling distance travelled by the tendon. The claw turns about 1 degrees per 4.3-5.0 microm of pulling distance travelled by the unguitractor. The arolium turns forward and downward simultaneously with flexion of the claws. The kinematic chain of the arolium lacks real condylar joints except the joint at the base of the manubrium. Other components are tied by flexible transmissions of the membranous cuticle. The walking hornet rests on distal tarsomeres of extended tarsi. If the retractor tendon inside the tarsus is fixed, passive extension of the tarsomeres might be replaced by claw flexion. Tarsal chain rigidity, measured with the force tester, increased when the retractor tendon was tightened. Probably, pull to the tendon compresses the tarsomeres, increasing friction within contacting areas of rippled surfaces surrounding condyles within articulations.     

Mating frequency and genetic relatedness of workers in the hornet Vespa analis (Hymenoptera: Vespidae)

Mating frequency of Vespa analis queens and the genetic relatedness of their workers was analyzed by DNA microsatellite genotyping. Of 20 colonies studied, 18 had a queen inseminated by a single male and two had queens each inseminated by two males. The estimated effective number of matings was 1.05 ± 0.037 (mean ± SE), with 75–85% of the offspring of the two multiply mated queens sired by a single male. The pedigree relatedness between nestmate workers averaged over the 20 colonies was estimated to be 0.74 ± 0.008, almost identical to the predicted value of 0.75 for colonies headed by a singly mated queen. Multiple matrilines; that is, the presence of workers not related to the current queens, were detected in six colonies, suggesting that queen replacement occurred via usurpation of the founding queens in these six colonies. These results demonstrate that the kin structure of V. analis is similar to that reported in other vespid species.     

Electrical properties of the cuticle,silk caps and comb of Oriental hornet Vespa orientalis (Hymenoptera : Vespidae)

Electrical and physical phenomena have been recorded and measured in the cuticle, silk caps and comb of the Oriental hornet Vespa orientalis (Hymenoptera : Vespdnae). Cuticle of active or narcotized live hornets as well as dead ones, produce, at optimal temperature for Vespinae biological activity, voltages of several hundred mV, currents of up to several hundred nA and the appropriate electric power. The cuticle has a large electrical capacitance, relative to its volume and contains non-linear and active electrical elements. A theoretical model was proposed to explain the capacitance phenomenon. An additional phenomenon observed is the production of electric energy under the influence of light and heat. Some electrical phenomena, especially the photoconductivity were measured also in 3 ant species.Measurements of the electrical capacitance of silk caps revealed that it is dependent on: (a) age of the pupa; immediately on pupation, the values are highest and diminish with maturation; (b) caste; capacitance for the queen pupae is 20–50 mF; and which is higher than for worker pupae, where it ranges between about 5–7 mF; and (c) location of the measuring electrodes; in the case of external-internal measurements, the values obtained were greater by 2 orders of magnitude than those obtained with both electrodes placed on the same side of the silk cap. In all cases, it was found that the capacitant values are high when compared with the size of the caps and the available commercial capacitors of the same size.The hornet comb may be regarded as comprised of an array of 3-dimensional capacitors linked in parallel, thereby forming a large dry battery having one negative pole — the pedicel — which grounds the comb, and one positive pole — the silk domes of the comb cells. The possibility that the electric energy stored in the comb cell walls may have a thermoregulatory function, serving both the brood and the adult nest population was discussed. We assumed that this mechanism is common for the combs of all social as well as many solitary wasps.     

Post‐hibernation ovary development in queens of the Japanese giant hornet Vespa mandarinia (Hymenoptera: Vespidae)

Ovary contents and insemination status were examined in overwintered queens of Vespa mandarinia that were collected using bait traps in Ibaraki Prefecture, eastern Japan. Catches in the traps occurred from late April to early July in 1999, though they were few or rather sporadic in and after June. Fertilization rates were 100% in 1999, and 93% in 1998 when a preliminary sampling was made. While the ovary index (number of eggs with size equal to or larger than adjacent nurse cells) of queens caught on the first few collection dates was small, it rapidly increased up to late May. Generalized linear model analyses showed that the ovary index depended both on collection date and the size (head width) of the queens. Mature eggs were almost absent in their ovaries before early May, whereas more than 70% of the queens had one or more mature eggs on and after May 10, suggesting the onset of nesting in early May.     

Complete mitochondrial genome of the black‐tailed hornet,Vespa ducalis (Hymenoptera: Vespidae): Genomic comparisons in Vespoidea

We sequenced the complete mitochondrial genome (mitogenome) of the black‐tailed hornet, Vespa ducalis (Hymenoptera: Vespidae). The genome was 15,779‐bp long and contained typical sets of genes [13 protein‐coding genes (PCGs), 22 tRNAs, and 2 rRNAs]. The V. ducalis A + T‐rich region was 166‐bp long and was the shortest of all sequenced Vespoidea genomes, including Vespa. The genome was highly biased toward A/T nucleotides—80.1 % in the whole genome, 77.8 % in PCGs, 83.4–85.6 % in RNAs, and 92.8 % in the A + T‐rich region. These values are well within the typical range for genes and regions of Vespoidea mitogenomes. Start and stop codons in several Vespa species—including V. ducalis—were diversified, despite these species belonging to the same genus. In comparison with the ancestral mitogenomes, Vespa mitogenomes—including that of V. ducalis—showed substantial gene rearrangement; however, we detected no gene rearrangement among Vespa species. We conducted phylogenetic reconstruction based on concatenated sequences of 13 PCGs and two rRNAs (12,755 bp ) in available species of Vespoidea—21 species in six subfamilies in two families (Vespidae and Formicidae). The Bayesian inference and maximum likelihood (ML) methods revealed that each family formed strong monophyletic groups [Bayesian posterior probability (BPP) = 1; ML, 100 %]. Moreover, V. ducalis and V. mandarinia formed a strong sister group (BPP = 1; ML, 94 %).     

Weight changes in adult hornets,Vespa affinis (Hymenoptera: Vespidae)

Summary After sexuals emerge from their cells, they remain within the nest for 8–11 days (males) or 13–14 days (queens). During this time their weight increases (males 38 %, queens 40 %) due to the laying down of fat in the gaster. The weight of workers does not change over the same period. The fat is utilized by the queen during the 4–5 month overwintering period. When queens emerge in spring their weight is at its lowest, and they feed on nectar. Ovarian development, which was delayed during winter, now begins. During summer, queens again gain weight but this is now due to the development of the ovaries. The queen lives for about 1 year.     

Workers and queens of the European hornetVespa crabro L. have colony-specific cuticular hydrocarbon profiles (Hymenoptera: Vespidae)

Summary Cuticular hydrocarbons were extracted from sixty individual workers from six colonies ofVespa crabro L. and analyzed by combined gas-chromatography/mass spectrometry. Discriminant analysis of the cuticular hydrocarbon profiles of workers and queens showed that the wasps could be grouped by colony and by caste. Stepwise discriminant analysis selected the components which were weighted most heavily in these analyses. Different combinations of cuticular hydrocarbons were important in grouping workers by colony, queens and workers by colony, and workers and queens by caste.     

Early collapse of Vespa simillima (Hymenoptera, Vespidae) colonies in central Japan

During the past 15 years in the Ina Valley, Japan, mature fifth instar larvae of the yellow hornet, Vespa simillima, have been occasionally ejected from colonies during September. During 2005, this unusual behavior was particularly widespread, and collection of several V. simillima colonies confirmed that very few fifth instar larvae were present. When compared with an average colony, constructed from 41 colonies collected 20 years previously, colonies in 2005 had 80% fewer fifth instar larvae, despite queen egg‐laying rates remaining similar. It may be that the amount of food provided to the larvae by workers is less than the amount of larval secretions received by workers, which is causing the larvae to become emaciated and preventing them from pupating. This phenomenon normally occurs naturally at the end of the colony cycle in November, when prey abundance decreases and larval secretions or other sources of carbohydrates, for example from ripe fruit, tree sap, or aphid secretions, are needed by sexuals to build up fat reserves. However, another possibility is that the unexplained appearance of this phenomenon in September, which is causing colonies to collapse without producing any or very few sexuals, is due to foragers feeding the larvae prey that are contaminated with insect growth regulators, via pesticides, which are known to prevent successful pupation.     

Occurrence of Vespa velutina Lepeletier from Korea, and a revised key for Korean Vespa species (Hymenoptera: Vespidae)

This is the first report of occurrence of Vespa velutina Lepeletier from Korea. The diagnosis and taxonomic relationship between geographical subspecies is provided, with a revised key to complement the previous taxonomic information of Korean Vespa species. The ecological aspect of the species is also discussed.     

Tracing the invasion characteristics of the yellow-legged hornet,Vespa velutina nigrithorax (Hymenoptera: Vespidae), in Korea using newly detected variable mitochondrial DNA sequences

The yellow-legged hornet, Vespa velutina nigrithorax (Hymenoptera: Vespidae), invaded South Korea in 2003 through Busan metropolitan city, which is located in the southeast region of the country. Previous studies aiming to trace the origin of V. velutina in Korea used a portion of mitochondrial (mt) COI and detected a single haplotype common to the site of origin. However, no subsequent study on invasive dynamics such as additional entry and/or another site of entry has been performed. In this study, segments of mt COI, CytB, and lrRNA were sequenced from 238 individuals collected in 11 Korean and two Japanese localities, but no variation in each gene was observed. Thus, we developed two intergenic spacer (IGS) sequences from the publicly available mt genome of V. velutina, which provided substantially increased variability (i.e., 19 haplotypes with 1.74% maximum sequence divergence in 1,129–1,146-bp-long concatenated sequences). Population genetic analyses using the concatenated sequences unexpectedly provided higher genetic diversity estimates in the northwest and southwest regions, both of which also harbor international cargo ports, than in the southeast region, in which Busan is located. Furthermore, this genetic result was roughly concordant with our questionnaire survey demonstrating that V. velutina was observed in apiaries located in the northwest and southwest regions up to 2012, when there was no reported prevalent distribution of the hornet beyond the southeast region. These results collectively suggest that the northwest and southwest regions of Korea are additional sites of V. velutina entry to the country, independent from the southeast region origin.     

Smaller nests of the hornet Vespa analis (Hymenoptera, Vespidae) are more severely affected by the strepsipteran parasite Xenos moutoni (Strepsiptera, Stylopidae) than are larger nests

The parasite Xenos moutoni inactivates workers and castrates reproductives of the Japanese hornet Vespa analis. Using over 500 nests, we investigated the relationship between hornet nest size (number of cells) and the proportion of parasitized adults (i.e., prevalence) in central Japan. Over 3 years, 36–48% of nests had more than one parasitized adult, and approximately 3–5% of female adults (mostly workers) and 1–3% of male adults were parasitized within the population. The number of cells did not differ between nests with and without parasitized workers; therefore, the effects of parasite-caused losses within the worker force on nest size appear to be negligible. However, the prevalence of parasitized workers tended to decrease with an increase in the number of cells. Our analysis indicated that the number of parasitized workers was consistently low and did not increase with nest size, resulting in a relatively high prevalence in small nests.     

Enzymatic and chemical properties of an endopeptidase from the larva of the hornet Vespa crabro.

An endopeptidase from the larvae of the hornet Vespa crabro has been purified to homogeneity. The enzyme has been characterized with respect to molecular weight, amino acid compositon, and amino- and carboxyl-terminal sequences. The catalytic properties of the hornet protease are similar to those of bovine chymotrypsin with respect to inactivation by phenylmethanesulfonyl fluoride and carbobenzoxyphenylalanine chloro ketone and preferential peptide bond cleavage at aromatic amino acid residues. In contrast to bovine chymotrypsin, the hornet protease is not inhibited by the basic pancreatic Kunitz inhibitor, soybean inhibitor, or chicken ovomucoid. The molecular weight, as determined by several independent methods, was found to be 14 500. The protease is a single-chain protein containing two disulfide bonds. The terminal sequences are: NH2-Ile-Val-Gly-Gly-Ile-Asp.....Gly-Lys-Tyr-Pro-Tyr-Gln-Val-Ser-Leu-Arg-COOH.     

Sphaerularia vespae sp. nov. (Nematoda, Tylenchomorpha, Sphaerularioidea), an endoparasite of a common Japanese hornet, Vespa simillima Smith (Insecta, Hymenoptera, Vespidae)

Sphaerularia vespae sp. nov., an endoparasite of a common Japanese hornet, Vespa simillima is described from Hokkaido, Japan, and its molecular sequence profiles are given. This newly discovered nematode appears to belong to the genus Sphaerularia, judging from its characteristic parasitic form, the uterium, which looks like a sausage with many verrucae on its surface. Sphaerularia vespae sp. nov. is distinguished from S. bombi, the only other nominal species of the genus, by the morphology of the male bursa, female tail, and anterior end of parasitic juveniles. SSU, ITS, and D2D3 (LSU) DNA sequences were determined for S. vespae sp. nov. and compared with those from related nematodes obtained from the GenBank database. The sequences from S. vespae sp. nov. were close to those of S. bombi and several another tylenchid entomoparasitic nematodes. Sphaerularia vespae sp. nov. parasitizes overwintering gynes of the hornet and practically sterilizes them, as S. bombi does for bumblebee gynes.     

Sperm structure and ultrastructure in the Hymenoptera (Insecta)

A light and electron microscopical survey of spermatozoan gross morphology and ultrastructure in the Hymenoptera is presented. Details are provided for the first time for members of the families Xyelidae, Argidae, Tenthredinidae, Diprionidae, Cephidae, Figitidae, Proctotrupidae, Diaprii- dae, Heloridae, Eurytomidae, Leucospidae, Perilampidae, Torymidae, Braconidae, Dryinidae, Sphecidae, Pompilidae and Vespidae. Spermatozoan length ranged from 8 μ m in some Braconidae to 500 μm in one chalcidoid. Considerable variation in gross morphology and ultrastructure were observed between taxa. Several phylogenetically informative characters were noted. Very small spermatozoa characterized most of the non-cyclostome subfamilies of Braconidae; spirally twisted axoneme and mitochondrial derivatives occur in the Eulophidae, Eurytomidae and Pteromalidae; spermatozoa with virtually indistinguishable head (nucleus and acrosome) regions characterized the Vespinae and Polistinae. The presence of well-developed spermatodesmata in the vas deferens and seminal vesicle characterize the Symphyta and were largely absent from other groups though they are occasionally present in some bees.