Tuberaphis owadai (Homoptera), a new aphid species forming a large gall on Styrax tonkinensis in northern Vietnam

Tuberaphis owadai sp. nov., an aphid species forming coral‐shaped galls on Styrax tonkinensis in northern Vietnam, is described. We found that the species produces many sterile second‐instar soldiers in the gall. The colony size of a large gall was estimated to be 178 000, approximately half of which were soldiers. Alates emerging from galls contained sexual embryos, indicating that the life cycle is monoecious (non‐host‐alternating). Predaceous larvae of the pyralid moth Assara seminivalis were found in several galls.     

Secondary host generation of the gall aphid Cerataphis jamuritsu (Homoptera)

Colonies of a Cerataphis species with well‐developed horns were found on the rattan Calamus quinquesstinervis in southern Taiwan. The morphology of first instar nymphs from the colonies accorded well with the morphology of first instar nymphs laid by alates of Cerataphis jamuritsu from galls on Styrax suberifolia, indicating that the rattan aphids are the secondary host generation of C. jamuritsu. Although the aphid colonies were attended by ants, the sharp horns of the first instar nymphs suggest that they might attack predators.     

Two closely related species of Nipponaphis (Hemiptera: Aphididae) that migrate between Distylium racemosum and Machilus trees in Japan

Nipponaphis loochooensis Sorin, 1996 and N. machilicola (Shinji, 1941) form fig‐shaped or globular galls on Distylium racemosum in southern Japan. Nipponaphis machilicola migrates to the lauraceous evergreen Machilus thunbergii. Nipponaphis loochooensis has also been supposed to migrate to M. thunbergii, but its secondary‐host generation has not been found in the field to date. Through sampling Nipponaphis aphids from trees of M. japonica in addition to M. thunbergii, and sequencing their mitochondrial DNA, we found that the two species form colonies on twigs of both Machilus species, and that the two at times colonize on the same trees and even form mixed colonies. Only N. loochooensis forms colonies on leaves of M. japonica, but neither species colonizes on leaves of M. thunbergii. The secondary‐host generations of the two species could be clearly discriminated from each other, based on morphology. It was confirmed by examining the type specimens of Nipponaphis amamiana Takahashi, 1962 that the name is a junior synonym of N. machilicola.     

Astegopteryx spinocephala (Hemiptera: Aphididae), a new aphid species producing sterile soldiers that guard eggs laid in their gall

Astegopteryx spinocephala sp. nov., a soldier‐producing aphid species forming banana‐bunch shaped galls on Styrax benzoides in northern Thailand, is described. We found that galls of the species are formed in approximately June and last for almost 1 year, and that the aphid completes its life cycle without migrating to secondary hosts. Many alate sexuparae appeared in March/April, when many sexuals (arostrate males and rostrate females) and eggs were found in live subgalls guarded by soldiers that plugged the ostiole with their sclerotized, spiny heads. Two healthy galls sampled in April contained 2799 and 2659 eggs, respectively. Many live galls were still found in May, and some at the beginning of June. These galls contained both active soldiers and eggs, some of which had already hatched. This indicates that soldiers of A. spinocephala guard eggs until they hatch in at least some galls.     

Defensive Behavior in Primary- and Secondary-Host Generations of the Soldier-Producing Aphid, Pemphigus bursarius (Hemiptera: Aphididae)

The genus Pemphigus comprises several species that produce soldiers (defensive morphs) in galls on the primary host. At the moment, it is unclear if host-alternating species also produce defenders on their secondary host. We therefore examined how P. bursarius morphs of the secondary host generations respond to predators to test whether they show defensive behavior. We further examined how this response compares with the antipredator behavior of soldiers in P. bursarius and P. spyrothecae occurring on the primary host. We performed two manipulative experiments using two predatory species to quantify the behavior of the different morphs in response to predators. In both experiments, secondary host morphs of P. bursarius showed no attacking behavior and antipredator behavior in these morphs was limited to escaping natural enemies by walking away. In contrast, the first instars of the primary host generations in both Pemphigus species showed attacking behavior and were capable of killing the predators.     

Description and ecological traits of a new species of Pitydiplosis (Diptera: Cecidomyiidae) that induces leaf galls on Pueraria (Fabaceae) in East Asia,with a possible diversification scenario of intraspecific groups

A gall midge that induces thick lenticular galls on leaflets of Pueraria species (Fabaceae) in Japan, mainland China, Taiwan and South Korea is described as Pitydiplosis puerariae sp. nov. (Diptera: Cecidomyiidae). Tanaostigmodes puerariae (Hymenoptera: Tanaostigmatidae), described earlier from mainland China as an inducer of the lenticular gall, is regarded to be an inquiline. Pitydiplosis puerariae is distinguishable from the only known congener, the Nearctic Pitydiplosis packardi, by the male genitalia with entire aedeagus and with hypoproct that is as long as cerci and bilobed with a U‐shaped emargination. DNA sequencing data indicate the existence of three genetically different intraspecific groups: (i) “YNT‐montana group” induces galls on Pueraria montana on the Yaeyama Islands, Japan and in northern Taiwan; (ii) “CT‐montana group” on P. montana in central Taiwan; (iii) and “JCK‐lobata group” on Pueraria lobata in mainland China, South Korea and Japan north of Okinoerabu Island. A possible diversification scenario of the three groups is hypothesized based on DNA sequencing data and geohistorical information. A distribution gap of the gall midge on five islands between Tokunoshima and Ishigaki Islands, Japan was confirmed by intensive field surveys. Ecological traits and adult behavior of Pity. puerariae are also described. Its possibility as a potential biological control agent against P. lobata seems counter‐indicated.     

Nipponaphis species (Aphididae: Hormaphidinae) that form green galls on Distylium racemosum in Japan

Some species of the aphid genus Nipponaphis (Nipponaphidini) form green, globular or fig‐shaped galls on the evergreen Distylium racemosum, their primary host. Molecular phylogenetic analyses of aphid samples collected from both their galls and secondary hosts indicated the occurrence of four species in Japan: N. distychii, N. distyliicola, N. loochooensis and N. machilicola. The four species could also be discriminated from one another in morphology. The name N. litseae turned out to be a junior synonym of N. distychii. Galls formed by N. machilicola are reported for the first time in this paper. The life cycles of the four species are briefly reviewed.     

Life history strategy and taxonomic position of gall midges (Diptera: Cecidomyiidae) inducing leaf galls on Styrax japonicus (Styracaceae)

Four gall midge species (Diptera: Cecidomyiidae) that induce leaf galls on Styrax japonicus (Styracaceae) were identified to generic level based on larval morphology. Three of these gall midges, which induce whitish hemiglobular galls, flattened subglobular galls, and purple globular galls, respectively, were identified as three genetically distinct species of Contarinia, and the remaining species, which induces globular galls with dense whitish hairs, was identified as a species of Dasineura. Field surveys in Fukuoka, Japan, revealed that adults of these gall midges emerged and oviposited in late March to mid‐April at Mount Tachibana (approximately 200 m a.s.l.) and in late April to early May at Mount Sefuri (about 1050 m a.s.l.), coinciding with the leaf‐opening season of S. japonicus. Larvae of these gall midges mostly developed into third instars by June and then left their galls and dropped to the ground. These species therefore have a life history strategy that differs from that of another S. japonicus‐associated gall midge, Oxycephalomyia styraci, which overwinters as the first instar in ovate swellings, matures rapidly in spring, and emerges directly from the galls.     

Formation of long‐lasting galls by overwintered nymphs in the Japanese aphid Quadrartus yoshinomiyai (Hemiptera: Aphididae: Hormaphidinae)

This paper examines the life history of a generation of galls created by the aphid Quadrartus yoshinomiyai (Hormaphidinae: Nipponaphidini) on its primary host plant, Distylium racemosum. First‐instar fundatrix nymphs of Q. yoshinomiyai initiated galls on stems of developing shoots in early April and incipient enclosed galls were found from later the same month. The galls lasted for up to 14 months, during which they grew to maturity, opened in early or mid‐April of the following year and dried up by the end of June. First‐instar fundatrix nymphs were found on winter buds, indicating that they hatched from eggs in autumn and overwintered as nymphs. These results suggest that Q. yoshinomiyai has a three‐year life cycle.     

Polymorphism of axillary bud galls induced by Rhopalomyia longitubifex (Diptera: Cecidomyiidae) on Artemisia princeps and A. montana (Asteraceae) in Japan and Korea, with designation of new synonyms

In the past, Rhopalomyia longitubifex, Rhopalomyia shinjii, and Rhopalomyia sp. (Diptera: Cecidomyiidae) have been regarded as independent species based on differences in the sizes and shapes of axillary bud galls induced on Artemisia montana (Asteraceae) in Japan and A. princeps in Japan and Korea. However, comparison of morphological features and molecular sequencing data indicate that these Rhopalomyia gall midges are identical and that the differences in gall shape are polymorphisms, although the measurements of gall height and diameter overlap slightly. This finding suggests that although galls have frequently been regarded as extensions of the phenotype of a species, differences in gall shape may not always be reliable for identifying gall‐inducing cecidomyiids. The older name, R. longitubifex, is applied to these gall midges, and the names that were applied to this species on later occasions are revised or synonymized. The mature and immature stages of R. longitubifex are redescribed and information on the distribution, host range, and gall size of this species is provided. We also discuss the role of gall polymorphism in the early stages of speciation.     

Life history strategy and adult and larval behavior of Macrodiplosis selenis (Diptera: Cecidomyiidae), a species that induces leaf‐margin fold galls on deciduous Quercus (Fagaceae)

Life historical, behavioral and ecological traits of Macrodiplosis selenis, which induces leaf‐margin fold galls on Quercus serrata, Q. mongolica and Q. dentata (Fagaceae) in Japan and South Korea, were studied. Daily activity and larval development indicate that M. selenis is a diurnal and univoltine gall midge. In April, females lay their eggs both on upper and under surfaces of fresh leaves. The duration of the egg stage varies from 5 to 9 days, depending on daily temperatures. Hatched larvae crawl to the upper surface of the leaf margin, where they start to induce galls. Larvae become full‐grown in October, drop to the ground in November and overwinter in cocoons on the ground, while larvae of congeners mature in May and drop to the ground in June. A relatively long period of the second larval stadium from July to October on the host trees seems to be effective for M. selenis in avoiding summer mortalities caused by predation and aridity on the ground and by ectoparasitoids that attack mature larvae or pupae on the host leaves. The spatial distribution pattern of M. selenis leaf galls is contagious and the mean gall density per leaf is significantly correlated with the mean crowding. This study adds new insights of life history strategy and adult and larval behavioral pattern to the ecological knowledge of gall midges, and these kinds of information are essential for further studies of M. selenis population dynamics and interactions with other Quercus‐associated herbivores.     

Life history characteristics and behaviour of the bamboo aphid,Pseudoregma bambucicola (Hemiptera: Pemphigidae), having sterile soldiers

Summary Life history and behavioural characteristics of the bamboo aphid,Pseudoregma bambucicola (Takahashi), which has sterile soldiers, were studied in the laboratory. The stadium of normal (fertile) first instar larvae was two times longer than that of second instar larvae, and the stadium of soldier-type (sterile) first instar larvae was much longer (max. 116 days) than the stadium of normal first instar, suggesting that soldiers are able to take nutrition from bamboo. Stimulation of larvae with breath, vibration of bamboo shoots or disturbing the larvae with the tip of a fine brush induced significantly more defensive acts by soldiers than those by normal larvae — the latter usually fled. Soldiers did not attack non-kin conspecific intruders or even aphids of different species, suggesting that, in this species, kin-recognition ability is low.     

Social parasitism and behavioral interactions between two gall-forming social aphids

We investigated ecological and behavioral aspects of the interactions between two social aphids, Pseudoregma bambucicola and Astegopteryx bambucifoliae (Hormaphidinae, Cerataphidini), both of which produce second-instar, sterile soldiers in galls formed on Styrax suberifolius, in Taiwan. By censusing their galls, either species was found to invade galls of the other species. Twenty-eight (58%) out of 48 A. bambucifoliae galls contained P. bambucicola, while four (6%) of 69 P. bambucicola galls contained A. bambucifoliae. Furthermore, P. bambucicola behaved like social parasites in galls of A. bambucifoliae. Colonies of P. bambucicola produced much fewer or even no soldiers compared with those in natal galls. Our experiments also revealed that individual aphids of P. bambucicola more successfully intruded into galls of A. bambucifoliae than into conspecific galls, and that guarding soldiers of P. bambucicola effectively prevented aphids from invading their galls and permitted only conspecific soldiers to join their colonies. These differences in behavior provide good explanations for the differences in the frequency of invaded galls found between the two species. Received 4 September 2007; revised 7 January 2008; accepted 17 January 2008.     

Spatial density‐dependent parasitism and specificity in the robber fly Mallophora ruficauda (Diptera: Asilidae)

The density‐dependence in parasitism by the robber fly Mallophora ruficauda (Diptera: Asilidae) on scarab beetle larvae (Coleoptera: Scarabaeidae) populations was studied in the present research. Mallophora ruficauda is a pestiferous species common in the open grasslands of the Pampas region of South America. Adults are predators of insects and larvae are solitary parasitoids of third instar larvae of several species of scarab beetle (Coleoptera: Scarabaeidae). In contrast with most studied host‐parasitoid interactions, host searching by M. ruficauda is carried out by both larvae and adults. Typically, robber fly females lay eggs on tall grasses from where larvae drop to the ground, and attack hosts which are buried in the soil. We carried out our study at two spatial scales close to 14 apiaries located in the provinces of Buenos Aires and Entre Ríos (Argentina). We found that parasitism is density‐independent at the larger spatial scale and inversely density‐dependent at the smaller one. We also found that M. ruficauda selects Cyclocephala signaticollis among several scarab beetle species. Specificity is observed both at large and small spatial scales. We discuss the implications of both host specificity and host searching behaviour on the observed parasitism patterns.     

Host preference and suitability of grasses for Oebalus pugnax

The rice stink bug, Oebalus pugnax (Fabricius) (Hemiptera: Pentatomidae: Carpocorini), though graminaceous, discriminates among its numerous host grass species. This could represent a feeding preference, it could be related to host suitability for growth and development. To clarify the role of host grass discrimination, two laboratory studies were conducted: (1) free‐choice tests to evaluate preferences of O. pugnax among 11 wild host grass species found in three rice‐producing counties of the central Mississippi Delta (MS, USA), and (2) no‐choice tests to evaluate the impact of rice (Oryza sativa L.), junglerice [Echinochloa colona (L.) Link], and dallisgrass (Paspalum dilatatum Poir.) (all Poaceae), on the development of O. pugnax from second instar to adult. In the free‐choice test, four experiments were conducted, each with four sets of host grass species and observed 1, 2, 4, 8, and 16 h after release in cages. Approximately 4 h was necessary for O. pugnax to settle on preferred host grasses. Oebalus pugnax showed a feeding preference for junglerice over all 10 other grass species. Bahiagrass, Paspalum notatum Flueggé, was the least preferred. The no‐choice tests showed significant effect of host grass species on O. pugnax mean development time of nymphal survival to adults. Survival of nymphs was lower and mean development time was longer on dallisgrass compared to rice and junglerice. Knowledge of O. pugnax rate of growth and development on host grasses could be useful in the future development of rice integrated pest management strategies.     

Behavioural and phylogeographic observations on Epipomponia nawai (Dyer, 1904): An East Asian moth (Lepidoptera; Epipyropidae) whose larvae are ectoparasitic on cicadas (Hemiptera; Cicadidae; Sonatini)

The lepidopteran family of the Epipyropidae contains currently 32 known species of moths with larvae that are parasitic on plant and leafhoppers as well as cicadas. This paper focuses on behavioural aspects of the final instar of the cicada parasite Epipomponia nawai (Dyer 1904) on the cicada Hyalessa maculaticollis (de Motschulsky 1866) and represents only the second report of the species from South Korea. This species is a new host for E. nawai in Korea, since in the past it had only been reported once from Meimuna opalifera and Oncotympana fuscata in Korea. Molecular relationships between specimens of E. nawai collected in China, Japan, and Korea were analysed. The Chinese haplotypes fell into two groups: one clearly related to the Korean haplotype, but the other tightly affiliated with the Japanese haplotype. The findings suggest two routes for the occurrence of E. nawai in mainland China. Once the host cicada was dead or too weak to move, the caterpillar crawled away from it, seeking a place to pupate. Very little is known about parasitic Lepidoptera and therefore any information on host species or behavioural aspects is of importance.     

Cryptic diversity of Melanochlamys sea slugs (Gastropoda,Aglajidae) in the North Pacific

North Pacific specimens of Melanochlamys sea slugs were examined morphologically (including the male reproductive morphology, shell and external coloration) and were sequenced for three genes (mitochondrial COI and 16S and nuclear H3). Phylogenetic and species delimitation analyses were used to determine the evolutionary relationships and species diversity among the specimens examined. Both molecular and morphological data revealed an unexpected high level of cryptic diversity. At least four distinct species occur on the Northwestern Pacific. Melanochlamys ezoensis occurs in Russia and temperate and cold areas in Japan. Three additional undescribed species occur in Japan and/or South Korea. One of the undescribed species occurs both in South Korea and in Japan, but only in Tokyo Bay, suggesting that it could be non‐native in Japan. Two distinct species occur on the Northeastern Pacific coast; Melanochlamys diomedea is widespread from Southern California to Alaska, whereas M. ezoensis was found only in San Francisco Bay, suggesting a human‐mediated introduction. This is further supported by the absence of records of M. ezoensis in San Francisco prior to 2001. The species diversity of Melanochlamys in the Northwestern Pacific is much greater than in the Northeastern Pacific; it is hypothesized that differences in geographic and ocean current system complexity might account for different responses to glacial extinction and postglacial expansion.     

Genetic divergence of Platycerus hongwonpyoi (Coleoptera: Lucanidae) in South Korea

We examined the genetic divergence of Platycerus hongwonpyoi Imura & Choe, 1989 in South Korea using the nuclear wingless (Wg) gene, internal transcribed spacer (ITS) region and mitochondrial cytochrome oxidase subunit I (COI) gene. We found no variation in Wg or ITS. Based on COI, P. hongwonpyoi was split into four well defined and one weakly supported clades, which were inferred to have diverged 2.11–1.33 Ma. The Platycerus hongwonpyoi population size seems to have decreased during the past several tens of thousands of years. The divergence times of major clades of P. hongwonpyoi were comparable with those involved in the speciation of certain Japanese species. Frequent overlapping of different clades at the same sites suggests the occurrence of secondary gene flow following differentiation in South Korea. In conclusion, the genus Platycerus underwent strikingly different divergence patterns in South Korea compared with Japan according to the disparate topographies of these two geographical areas.     

Nine new species of Dasineura (Diptera: Cecidomyiidae) from flowers of Australian Acacia (Mimosaceae)

Abstract. Thirteen species of Australian acacias are invasive plants in agricultural and native vegetation areas of South Africa. Biological control programmes for Australian acacias in South Africa have been implemented and are aimed at suppressing reproductive vigour and, in some cases, vegetative growth of these weeds. Gall-forming midges are under consideration as potential biological control agents for invasive acacias in South Africa. Entomological surveys in southern Australia found a diverse cecidomyiid fauna associated with the buds, flowers and fruits of Acacia species. Nine new Dasineura species are described and two species, D. acaciaelongifoliae (Skuse) and D. dielsi Rübsaamen, are redescribed. The newly described taxa are D. fistulosa sp.n. , D. furcata sp.n. , D. glauca sp.n. , D. glomerata sp.n. , D. oldfieldii sp.n. , D. oshanesii sp.n. , D. pilifera sp.n. , D. rubiformis sp.n. and D. sulcata sp.n. All eleven species induce galls on ovaries and prevent the formation of fruit. Two general types of gall are caused. Type A comprises woody, tubular galls with larvae living inside ovaries (D. acaciaelongifoliae, D. dielsi, D. fistulosa, D. furcata, D. glauca, D. glomerata, D. oldfieldii). Type B includes soft-tissued, globose galls that belong to four subtypes: inflated, baglike, hairy galls with larvae living between ovaries (D. pilifera); pyriform, pubescent swellings with larvae living inside ovaries (D. rubiformis); globose, hairy, swellings with larvae living superficially on ovaries in ovoid chambers (D. oshanesii); and inconspicuous, glabrous swellings with larvae living superficially on ovaries in shallow groovelike chambers (D. sulcata). The gall types are associated with a particular pupation pattern. In type A galls, larvae pupate within larval chambers in galls, whereas in type B galls pupation takes place between ovaries in galls or in the soil beneath the host tree. Gall midges responsible for the same general gall type are morphologically related and differ from species causing the other gall type. Phylogenetic analysis of a 410 bp fragment of the mitochondrial cytochrome b gene supports the division of the gall midge species into two groups except for D. sulcata, which appears as a subgroup of the group causing type A galls. The interspecific divergence values in group A species were between 0.5 and 3.9% with intraspecific divergence estimates of 0–0.2%. Gall midges causing type B galls had interspecific divergence values of 4.6–7.3% and intraspecific divergence values of 0–3.7%. Closely related biology and morphology together with low cytochrome b divergence estimates suggest a more recent speciation in group A when compared with species of group B. Dasineura rubiformis and D. dielsi are proposed as potential biological control agents for Acacia mearnsii De Wild. and Acacia cyclops A. Cunn. ex G. Don, respectively, in South Africa due to their narrow host range and ability to form high population densities that reduce seed formation. Both species produce galls with low biomass, which makes them compatible with commercial exploitation of their host species in Africa.     

Larval descriptions of five Byctiscus species (Coleoptera: Rhynchitidae) in Korea

The genus Byctiscus reported in Korea in five of nine species (B. lacunipennis, B. rugosus, B. puberulus puberulus, B. princeps and B. coerulans) is described for the last instar larvae. In addition, the cradle structure and host‐plants are provided.