Shin-ichi Akimoto

Ant attendance changes the sugar composition of the honeydew of the drepanosiphid aphid Tuberculatus quercicola

Mutualistic interactions between aphids and ants are mediated by the honeydew produced by aphids. Previous work showed that when attended by the ant Formica yessensis, nymphs of the aphid Tuberculatus quercicola developed into significantly smaller adults with lower fecundity than when not ant-attended. This study tested the hypothesis that this cost of ant attendance arises through changes in the quality and quantity of honeydew. Ant-attended and ant-excluded aphid colonies were prepared in the field, and the sugar concentration and sugar composition of the honeydew of ant-attended colonies were compared with those of ant-excluded colonies. The frequency and amount of honeydew excretion were also quantified in the two types of colonies. The aphids excreted smaller droplets of honeydew more frequently in ant-attended colonies than in ant-excluded colonies. There was no significant difference in total sugar concentration between the honeydew of ant-attended aphids and ant-excluded aphids. However, ant-attended aphids produced honeydew containing a significantly lower proportion of glucose and higher proportions of sucrose and trehalose than did ant-excluded aphids. These results suggest that the enhanced rate of honeydew-excretion behavior under ant attendance led to changes in the aphids physiological status. We suggest that the increase in the proportions of sucrose and trehalose in honeydew leads to a shortage of carbohydrates available for energy metabolism, resulting in lower performance of the aphids under ant attendance.

Genetic differentiation as a result of adaptation to the phenologies of individual host trees in the galling aphid Kaltenbachiella japonica

Abstract. 1 The adaptation of the non‐migratory galling aphid Kaltenbachiella japonica (Matsumura) to the budburst phenologies of individual host trees (Ulmus davidiana var. japonica Nakai) was investigated. There was a large variation in budburst time between individual host trees. We tested a hypothesis that aphid populations on respective host trees are genetically differentiated through adaptation to the budburst phenologies of host trees. 2 There was significant correlation between the budburst time of eight host trees in two successive years. 3 Tree‐associated populations differed significantly in egg‐hatching time, and the mean hatching time was significantly correlated with the mean budburst time of respective host trees. 4 Mating was made between male and female sexuales of the same clone (i.e. self‐mating) to obtain selfed eggs. Hatching patterns of selfed eggs showed that there was a substantial amount of genetic variance in hatching time between clones (galls). Nested ANOVA demonstrated that the between‐tree component contributed more to the phenotypic variance than the within‐tree or within‐gall components. 5 Reciprocal crosses between males and females of different clones furthermore demonstrated significant differences in hatching time between cross combinations, but no significant differences between the two reciprocals within combinations. This result suggests that there are no maternal effects for the timing of egg hatch and that the differences between selfed lines are attributable to genetic variance. 6 The fine‐scale adaptation hypothesis was supported by egg‐hatching experiments, which further suggest that budburst phenology varying between individual trees is acting as intense selective pressure on the egg‐hatching time of associated insects.

Flexibility in the composition and concentration of amino acids in honeydew of the drepanosiphid aphid Tuberculatus quercicola

Abstract 1. Mutualistic interactions between aphids and ants are mediated by honeydew that aphids produce. Previous work showed that when attended by the ant Formica yessensis Forel (Hymenoptera: Formicidae), nymphs of the aphid Tuberculatus quercicola (Matsumura) (Homoptera: Aphididae) developed into significantly smaller adults with lower fecundity than did nymphs that were not ant attended.

Taxonomic status of the Bemisia tabaci complex (Hemiptera: Aleyrodidae) and reassessment of the number of its constituent species.

Bemisia tabaci (Hemiptera: Aleyrodidae) is one of the most important insect pests in the world. In the present study, the taxonomic status of B. tabaci and the number of species composing the B. tabaci complex were determined based on 1059 COI sequences of B. tabaci and 509 COI sequences of 153 hemipteran species. The genetic divergence within B. tabaci was conspicuously higher (on average, 11.1%) than interspecific genetic divergence within the respective genera of the 153 species (on average, 6.5%). This result indicates that B. tabaci is composed of multiple species that may belong to different genera or subfamilies. A phylogenetic tree constructed based on 212 COI sequences without duplications revealed that the B. tabaci complex is composed of a total of 31 putative species, including a new species, JpL. However, genetic divergence within six species (Asia II 1, Asia II 7, Australia, Mediterranean, New World, and Sub Saharan Africa 1) was higher than 3.5%, which has been used as a threshold of species boundaries within the B. tabaci complex. These results suggest that it is necessary to increase the threshold for species boundaries up to 4% to distinguish the constituent species in the B. tabaci complex.

Allometric Patterns of Heads and Genitalia in the Stag Beetle Lucanus maculifemoratus (Coleoptera: Lucanidae)

Abstract Males of Lucanus maculifemoratus Motschulsky are markedly variable in size of body and heads. In this study, we investigated the possible causes that affect the development of the male heads and genitalia by comparing variability of weight among body parts and allometry in each part to body size. Genitalia varied least in weight among males, and the frequency distribution of their weight was approximately normal. In contrast, the frequency distribution of head weight exhibited a conspicuous skew. Significant positive allometry was found for heads, whereas genitalia showed significant negative allometry. Heads that are used for fighting are allometrically highly variable, whereas genitalia are highly stable, suggesting variable strategies for obtaining mates while maintaining equally sexually functional genitalia. The low variability and low allometric coefficient with body size for genitalia may indicate that the development mechanism for genitalia is separated from that for other body parts, as a result of sexual selection.

Aphid galls accumulate high concentrations of amino acids: a support for the nutrition hypothesis for gall formation

The nutrition hypothesis for the adaptive significance of insect gall formation postulates that galls accumulate higher concentrations of nutritive compounds than uninfested plant tissue, resulting in a high performance of the gall former. This hypothesis has been supported by some taxa of gall insects, but not by taxa such as cynipid wasps. Aphid galls are expected to require higher levels of nitrogen than other insects’ galls with a single inhabitant, because aphid galls are required to sustain a number of aphids reproducing parthenogenetically over two generations. The present study tested this hypothesis by evaluating aphid performance and amino acid concentration in phloem sap, using the aphid Rhopalosiphum insertum (Walker) (Homoptera: Aphididae), which establishes colonies on leaves of Sorbus commixta Hedlund or in galls of the aphid Sorbaphis chaetosiphon Shaposhnikov (Homoptera: Aphididae). We prepared the gall and non‐gall treatments on trees of S. commixta, in which R. insertum fundatrices were reared and allowed to reproduce. In S. chaetosiphon galls, R. insertum colonies propagated more rapidly, and the second generation grew larger and more fecund than on ungalled leaves. The amount of amino acids exuding from cut galled leaves was fivefold that in ungalled leaves; however, there was no significant difference in the amino acid composition between galled and ungalled leaves. In the intact leaves, total amino acid concentration in the phloem sap declined rapidly from late April to late May; however, the galls retained this high amino acid concentration in developing leaves for 1 month. These results indicate that the improved performance in R. insertum is ascribed to the increased concentration of amino acids in galled leaves. We suggest that S. chaetosiphon galls function to promote the breakdown of leaf protein, leading to an increased performance of gall‐inhabiting aphids.

Local adaptation and host race formation of a gall-forming aphid in relation to environmental heterogeneity

SummaryThe process of host race formation in the aphid Tetraneura yezoensis is examined in relation to its population structure. T. yezoensis induces pouch galls on new leaves of Ulmus davidiana and U. laciniata. Its populations on the two host species are often sympatric. Fundatrices found on one elm species, when reciprocally transplanted to the other, suffered greatly reduced average fitness. This shows that aphid populations associated with the two elm species are genetically differentiated in physiological traits. Individual trees of each elm species showed large differences in susceptibility to gall formation and in bud burst time, and such between-tree variations were consistent over years. Overwintered eggs taken in early spring from four trees (two from each species) were incubated under the same temperature conditions. The average hatching time differed significantly even between populations from conspecific trees, and the sequence of egg hatching paralleled that of the leafing of those four trees. This between-tree difference in hatching time was consistent over years and was found to be genetic, showing that gene flow between aphid populations on separate trees is often restricted. The heterogeneity in host traits may have promoted the evolution of philopatry in this aphid. Of the fundatrices that hatched on a tree of one elm species, a few precent were preadapted to gall formation on the other elm species. This suggests that the formation of a new host race proceeds parapatrically under disruptive selection and at a low level of gene flow. Evidence was actually obtained that a small fraction of Tetraneura alates are passively transported and land on non-host plants.

Morphological abnormalities in gall-forming aphids in a radiation-contaminated area near Fukushima Daiichi: selective impact of fallout?

To evaluate the impact of fallout from the Fukushima Daiichi Nuclear Power Plant accident on organisms, this study compared the morphology and viability of gall-forming aphids between the Fukushima population and control populations from noncontaminated areas. This study, in particular, focused on the morphology of first-instar gall formers derived from the first sexual reproduction after the accident. Of 164 first instars from Tetraneura sorini galls collected 32 km from Fukushima Daiichi in spring 2012, 13.2% exhibited morphological abnormalities, including four conspicuously malformed individuals (2.4%). In contrast, in seven control areas, first instars with abnormal morphology accounted for 0.0–5.1% (on average, 3.8%). The proportions of abnormalities and mortality were significantly higher in Fukushima than in the control areas. Similarly, of 134 first instars from T. nigriabdominalis galls, 5.9% exhibited morphological abnormalities, with one highly malformed individual. However, of 543 second-generation larvae produced in T. sorini galls, only 0.37% had abnormalities, suggesting that abnormalities found in the first generation were not inherited by the next generation. Although investigation is limited to one study site, this result suggests that radioactive contamination had deleterious effects on embryogenesis in eggs deposited on the bark surface, but a negligible influence on the second generation produced in closed galls. Furthermore, analysis of both species samples collected in spring 2013 indicated that the viability and healthiness of the aphids were significantly improved compared to those in the 2012 samples. Thus, the results of this study suggest the possibility that a reduced level of radiation and/or selection for radiation tolerance may have led to the improved viability and healthiness of the Fukushima population.

Phenotypic selection on the process of gall formation of a Tetraneura aphid (Pemphigidae)

1. To clarify the evolutionary interactions between a galling aphid Tetraneura sp. and its host plant Ulmus davidiana var. japonica Nakai, we examined the pattern of phenotypic selection acting on the process of gall formation by multiple regression analyses and field experiments. 2. Two hypotheses have been advanced to explain variation in performance of gall insects within host plants. The «spatial heterogeneity hypothesis» postulates that size differences among leaves or shoots chosen by gall formers are primarily responsible for differential performance. The «synchronization hypothesis» emphasizes the effect of timing of insect attack relative to bud burst of the host plant as a source of the variation. 3. Multiple regression analysis showed that leaf size and shoot length did not contribute at all to fecundity in galls

Allelic and genotypic diversity in long-term asexual populations of the pea aphid, Acyrthosiphon pisum in comparison with sexual populations

Many aphid species exhibit geographical variation in the mode of reproduction that ranges from cyclical parthenogenesis with a sexual phase to obligate parthenogenesis (asexual reproduction). Theoretical studies predict that organisms reproducing asexually should maintain higher allelic diversity per locus but lower genotypic diversity than organisms reproducing sexually. To corroborate this hypothesis, we evaluated genotypic and allelic diversities in the sexual and asexual populations of the pea aphid, Acyrthosiphon pisum (Harris). Microsatellite analysis revealed that populations in central Japan are asexual, whereas populations in northern Japan are obligatorily sexual. No mixed populations were detected in our study sites. Phylogenetic analysis using microsatellite data and mitochondrial cytochrome oxidase subunit I (COI) gene sequences revealed a long history of asexuality in central Japan and negated the possibility of the recent origin of the asexual populations from the sexual populations. Asexual populations exhibited much lower genotypic diversity but higher allelic richness per locus than did sexual populations. Asexual populations consisted of a few predominant clones that were considerably differentiated from one another. Sexual populations on alfalfa, an exotic plant in Japan, were most closely related to asexual populations associated with Vicia sativa L. The alfalfa‐associated sexual populations harboured one COI haplotype that was included in the haplotype clade of the asexual populations. Available evidence suggests that the sexuality of the alfalfa‐associated populations has recently been restored through the northward migration and colonization of alfalfa by V. sativa‐associated lineages. Therefore, our results support the theoretical predictions and provide a new perspective on the origin of sexual populations.