The first record of genus Bruggmanniella (Diptera: Cecidomyiidae) from Taiwan with description of a new species inducing bud galls on Neolitsea parvigemma (Lauraceae)

The genus Bruggmanniella Tavares is newly discovered from Taiwan and Bruggmanniella brevipes sp. n. is described as new to science. This is the southernmost species of Bruggmanniella found in the Old World. Furthermore, the gall midge induces bud galls on Neolitsea parvigemma (Hayata) Kaneh (Lauraceae), an endemic species in Taiwan, and the plant genus Neolitsea is the third host genus of Lauraceae-associated Bruggmanniella in East Asia. The knowledge of its distribution and host information provide us to shed the light on evolutionary and biogeography issue of East Asian Bruggmanniella.www.zoobank.org/urn:lsid:zoobank.org:pub:820320EC-17BC-4E00-A401-2DBDA3EEE9C4     

Life history traits of Pseudasphondylia rokuharensis (Diptera: Cecidomyiidae) affecting emergence of adults and synchronization with host plant phenology

To identify adaptive strategies of gall-inducing cecidomyiids, we studied the life history traits of Pseudasphondylia rokuharensis (Diptera: Cecidomyiidae), a univoltine species inducing fruit galls on a deciduous shrub, Viburnum dilatatum. Although univoltine gall midges that are associated with trees or shrubs are usually difficult to rear from immature stages under artificial conditions, we successfully reared P. rokuharensis from first instar to adult. Mature larvae entered diapause in autumn, and the diapause terminated after low temperature treatment (4 mo of incubation at 5 degrees C). The adults emerged within a shorter period of time than those without low temperature treatment. Thus, the diapause provides better synchronization of adult emergence in the spring. Because adults live for only a few days, this synchronization improves the chances of adults finding a mate. This is the first report on the mechanism of larval diapause termination for univoltine and tree- or shrub-associated gall midges. The numbers of days needed for adults to emerge under laboratory conditions after low temperature incubation were significantly different between two different localities. This result might suggest the possible existence of ecoclines in their life history parameters.     

Adult behavior of an ambrosia gall midge Illiciomyia yukawai (Diptera: Cecidomyiidae) and synchronization between its emergence and host plant phenology

The adult behavior of an ambrosia gall midge Illiciomyia yukawai (Diptera: Cecidomyiidae) that induces leaf galls on Illicium anisatum (Illiciaceae) was studied at the population level from 1977 to 1995 in Kagoshima, Japan. Most males emerged between 0:00 and 08:00 h and females between 05:00 and 11:00 h. Males swarmed around the host trees between 05:00 and 11:00 h. Mating occurred on the host leaves mainly between 06:00 and 08:00 h. Females then left the host trees for somewhere else, possibly to collect symbiont fungal conidia. From 08:00 to 16:00 h, females were observed ovipositing into the host shoots. The low development threshold temperature for overwintered larvae was 14°C, while the thermal constant for emergence differed with individuals. Thermal totals above 14°C up to the 50% emergence date varied yearly from 33.1 to 68.7 degree‐days. The 50% emergence date varied from 9 to 18 May. The thermal totals significantly correlated with the 50% emergence date but did not correlate with the date when 50% of shoots became suitable for oviposition. Thus, the host‐plant responded to thermal effects differently from the gall midge. Illiciomyia yukawai has been synchronizing well with the host‐plant phenology but will suffer from asynchrony when global warming becomes more conspicuous.     

Insect galls on Bauhinia cupulata (Fabaceae): morphotypes characterization and description of a new species of Schizomyia (Cecidomyiidae,Diptera)

Galls on Bauhinia cupulata (Fabaceae) were investigated in two physiognomies of the Cerrado, riparian and dry forest, in the municipality of Barreiras (Western Bahia, Brazil) from October 2012 to July 2014. Four insect gall morphotypes were found. They were characterized based on shape, color, indumentum, plant organ of occurrence and galling taxon. A new galling species, Schizomyia barreirensis, is described, illustrated (larva, pupa, male, female and gall) and compared to other Neotropical congeneric species.     

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.     

Studies on the distribution pattern of the pine needle gall midge,Thecodiplosis japonensis Uchida etInouye (Diptera: Cecidomyiidae) in a pine forest

Numerical changes and distribution patterns of the pine needle gall midge, Thecodiplosis japonensisUchida etInouye , were studied during the period from 1978 to 1979 in a young plantation of Pinus thunbergii in Shiga Prefecture, Japan. The survivorship curve of this species was characterized by a low mortality of larvae in galls and two high mortalities before the formation of galls and during the overwintering period in soil. The within and between-trees distributions of eggs and larvae in galls were examined by using the regression method. The egg distribution per shoot was aggregative both within and between host plants. The within-tree variations in numbers of eggs per shoot were related to the differences in the abundance of available needles for oviposition per shoot among the canopy layers. The between-tree variations reflected the heterogeneous emergence of adult females in the study plot. The degree of aggregation increased from egg to gall stage in both within- and between-tree distributions and the increase was explained by the different mortality of larvae within trees and the inversely density-dependent mortality between trees. The distribution patterns in the soil habitat stages were examined by the patchness index (). This species showed aggregative distributions in soil stages. There was a correlation in spatial patterns of adult emergence between the successive generations. The distribution properties of this species were discussed in connection with the population dynamics and the availability of host plants in the study plot.     

In situ occurrence of a gall midge (Insecta,Diptera, Cecidomyiidae) on fossilized angiosperm leaf cuticle fragments from the Pliocene sediments of eastern India

In situ preservation of fossil insect damage in plant fossils is an excellent tool to study the coevolution of flora and fauna through geological time, but finding both damage and the insect causing that damage in the same specimen is a very rare phenomenon. Galling is a common form of angiosperm leaf damage, which can be regarded as a kind of extended phenotype of the causal insects, essentially the gall midges, but galls usually lack remains of the insects themselves. Here we report the in situ occurrence of a gall midge (Insecta, Diptera, Cecidomyiidae) as well as its pupal exuviae on the abaxial cuticular surface of fossilized leaf cuticle fragments of Fabaceae leaves (cf. Albizia) that also bear galls, recovered from the latest Neogene (Rajdanda Formation, Pliocene) sediments of the Chotonagpur Plateau, Jharkhand, northeastern India. This Pliocene gall midge features well-preserved legs, segmented antenna with distinct and enlarged scape, elongate curved setae, and longer than broad terminal plate of the ovipositor lamellae. The in situ presence of a gall midge on a host fabaceous leaf cuticle indicates the existence of a host-ectoparasite relationship in the ancient warm and humid tropical monsoon-influenced forests of eastern India during the Pliocene. This is the first authentic fossil record of an in situ phytophagous insect of Cecidomyiidae from India, as well as southeast Asia. Although the identification of the recovered phytophagous insect associated with the fossil leaf cuticle is only possible to family level, this find reveals that such plant-insect relationships existed in the Pliocene of eastern India.     

Structure of floral galls of Byrsonima sericea (Malpighiaceae) induced by Bruggmanniella byrsonimae (Cecidomyiidae,Diptera) and their effects on host plants

Galls are anomalies in plant development from parasitic origin, and affect cellular differentiation or growth of plants. This parasite–plant interaction occurs in many environments and typically in vegetative organs of plants. The existence of galls in reproductive organs and their effects on the host plant are seldom described in the literature. In this paper, we present a novel study of galls in plants of the neotropical region. Galls of Bruggmmaniella byrsonimae develop in the flower buds of Byrsonima sericea DC. (Malpighiaceae) and affect development of the reproductive organs and the reproductive effort of these plants. The sepals and petals show hypertrophy of parenchyma tissues after differentiation, and the stamens exhibit degeneration of the sporogenic tissue. The gynoecium is not entirely developed; ovary and ovules are often absent. Changes in vascular tissues are also frequent, which may indicate high demand for nutrient resources by the new tissues initiated by the larva. We compared the amount of inflorescences, galls and fruits to evaluate possible effects on host reproduction. The results suggest that the Cecidomyiidae galls in flower organs affect fruit set and the reproductive success of B. sericea.     

Description of a new Procystiphora species (Diptera: Cecidomyiidae) inducing galls in culms of Sasa nipponica (Poaceae) in Japan, and the first discovery of an inverted ovipositor in Diptera

Procystiphora uedai sp. nov., a cecidomyiid inducing subglobular galls on Sasa nipponica Makino and Shibata on Mount Ôdaigahara, Nara Prefecture, Japan is described herein. This species is distinguishable from the three known congeners by the following characteristics of the female post-abdomen: tergite VIII concave at both anterior and posterior margins; tergites VII, VIII and sternite VII unsclerotized; ovipositor–dorsoventrally inverted. Most individuals of this gall midge are univoltine and pupate in the galls in early September, but some individuals enter prolonged diapause at the third larval stadium and remain in the mature galls until the following year. Larvae of this species are attacked by two parasitoid species, Pediobius sasae Hansson (Eulophidae) and Torymus sp. (Torymidae).     

Effect of soil burial depth and wetting on mortality of diapausing larvae and patterns of post-diapause adult emergence of sorghum midge, Stenodiplosis sorghicola (Coquillett) (Diptera: Cecidomyiidae)

Abstract  In south-eastern Queensland, Australia, sorghum planted in early spring usually escapes sorghum midge, Stenodiplosis sorghicola , attack. Experiments were conducted to better understand the role of winter diapause in the population dynamics of this pest. Emergence patterns of adult midge from diapausing larvae on the soil surface and at various depths were investigated during spring to autumn of 1987/88–1989/90. From 1987/88 to 1989/90, 89%, 65% and 98% of adult emergence, respectively, occurred during November and December. Adult emergence from larvae diapausing on the soil surface was severely reduced due to high mortality attributed to surface soil temperatures in excess of 40°C, with much of this mortality occurring between mid-September and mid-October. Emergence of adults from the soil surface was considerably delayed in the 1988/89 season compared with larvae buried at 5 or 10 cm which had similar emergence patterns for all three seasons. In 1989/90, when a 1-cm-deep treatment was included, there was a 392% increase in adult emergence from this treatment compared with deeper treatments. Some diapausing larvae on the surface did not emerge at the end of summer in only 1 year (1989/90), when 28.0% of the larvae on the surface remained in diapause, whereas only 0.8% of the buried larvae remained in diapause. We conclude that the pattern of emergence explains why spring plantings of sorghum in south-eastern Queensland usually escape sorghum midge attack.     

A new Asphondylia species (Diptera: Cecidomyiidae) and a eulophid wasp (Hymenoptera) inducing similar galls on leaf buds of Schoepfia jasminodora (Schoepfiaceae), with reference to their ecological traits and a description of the new gall midge

Different gall inducers belonging to distinct insect orders are rarely known to induce similarly shaped galls on the same host plant organs. We report that Asphondylia tojoi Elsayed & Tokuda sp. nov. (Diptera: Cecidomyiidae) and Ceratoneura sp. (Hymenoptera: Eulophidae) induce galls on leaf buds of Schoepfia jasminodora Sieb. et Zucc. (Schoepfiaceae). We describe the gall midge species as new to science and report a phylogenetic analysis for known Japanese Asphondylia species. We also describe life histories of the two species, based on monthly surveys during 2015–2017: although both species are multivoltine, A. tojoi overwinters as first instars in galls, whereas Ceratoneura sp. possibly does so as adults outside the galls. In addition, the internal structure of galls differed between the two species. Galls containing A. tojoi consist of a single chamber with inner walls clearly covered with whitish fungal mycelia after the gall midges develop into second instars. Those containing the Ceratoneura sp. have multiple chambers with hard black inner walls. Although some eulophids are known to be inquilines of galls induced by Asphondylia species, we consider that the Ceratoneura sp. is probably a true gall inducer because of the different gall structure and absence of fungal mycelia in their galls. This is the first report detailing the annual life history of a Ceratoneura species. Asphondylia tojoi represents the first example of monophagous Asphondylia species with a multivoltine life history on a deciduous tree.     

Taxonomy and molecular phylogeny of Daphnephila gall midges (Diptera: Cecidomyiidae) inducing complex leaf galls on Lauraceae, with descriptions of five new species associated with Machilus thunbergii in Taiwan

Five new species of the genus Daphnephila (Diptera: Cecidomyiidae: Asphondyliini), D. ornithocephala, D. stenocalia, D. sueyenae, D. taiwanensis, and D. truncicola, all associated with Machilus thunbergii (Lauraceae), are described from Taiwan, and one previously known species, D. machilicola, is redescribed from Japan. Among the five new species, D. truncicola induces stem galls and the other four species induce leaf galls. A molecular phylogenetic analysis based on partial sequences of the mitochondrial cytochrome oxidase subunit I (COI) gene suggests that in this genus the stem-galling habit is a more ancestral state compared to the leaf-galling habit. Daphnephila seems to be of tropical origin and to have dispersed to Japan through Taiwan.     

Two new species of Torymus Dalman, 1820 (Hymenoptera: Torymidae) parasitizing Celticecis japonica Yukawa and Tsuda, 1987 (Diptera: Cecidomyiidae) that induce leaf galls on Celtis species (Ulmaceae) in Japan

Torymus celticolus Matsuo sp. nov. and Torymus celtidigalla Matsuo sp. nov. (Hymenoptera: Torymidae) are described as parasitoids of larvae of Celticecis japonica Yukawa & Tsuda, 1987 (Diptera: Cecidomyiidae) that induce leaf galls on Celtis species (Ulmaceae) in Japan. A key to species of the genus Torymus known in Japan is provided.     

Host plant manipulation by fig gall midges (Diptera: Cecidomyiidae) which induce emergence projections on flower galls in the syconia of Ficus microcarpa (Moraceae)

We observed that unique projections developed from female flower galls induced by three unidentified cecidomyiid species in the syconia of Ficus microcarpa (Moraceae) on Okinawa and Amami Islands, Japan. The three cecidomyiids (sp. 1, 2 and 3) were tentatively distinguished by the differences in the shape of the projections. The projection of sp. 1 started to develop from the bottom of each gall before emergence, broke the skin of the syconium, and developed finally up to 5–6 mm in length within 6–8 h. During this period, the pupa oriented its head towards the bottom of the gall. After the projection fully elongated, the pupa pushed open the bottom of the projection with its head. The projection was easily removed from the gall at the base. The pupa quickly crawled half way out of the gall through the opening at the bottom of the projection and an adult then emerged. The projection did not develop when other hymenopteran gall inhabitants emerged. The projection was derived from plant tissues consisting of a mass of small square cells in the basal and distal portions and regularly arranged long cells in the middle portion. No projection was induced by the application of gibberellin's paste to the bottom of syconia. The gall midge seemed to manipulate the fig plant to develop the projection before emergence, so that the pupa can easily pass through the sticky epidermis of the syconium. The emergence of sp. 2 and 3 could not be intensively observed.     

Phylogeny of the Baldratiina (Diptera: Cecidomyiidae) inferred from morphological, ecological and molecular data sources, and evolutionary patterns in plant-galler relationships

The phylogeny of the gall-midge subtribe Baldratiina (Diptera: Cecidomyiidae) was reconstructed from molecular (partial sequence of the mitochondrial 12S rDNA), morphological and ecological data sets, using 16 representative species of most of the genera. The morphological and ecological data were combined in a single character matrix and analyzed separately from the molecular data, resulting in an eco-morphological cladogram and a molecular cladogram. Attributes of galls and host associations were superimposed on the molecular cladogram in order to detect possible trends in the evolution of these traits. The cladograms resulting from the two independent analyses were statistically incongruent, although both provide evidence for the monophyly of the genera Baldratia and Careopalpis and the paraphyly of the genera Stefaniola and Izeniola. The results suggest a minor impact of the morphological characters traditionally used in the classification of the Baldratiina, whereas ecological data had a major impact on the phylogenetic inference. Mapping of gall and host attributes on the molecular cladogram suggests that multi-chambered stem galls constitute the ancestral state in the subtribe, with several subsequent shifts to leaf galls. It is concluded that in contrast to other studied groups of gall insects, related baldratiine species induce different types of galls, attesting to speciation driven by gall-type shifts at least as often as host shifts.     

Long term oviposition preference and larval performance of Schizomyia macrocapillata (Diptera: Cecidomyiidae) on larger shoots of its host plant Bauhinia brevipes (Fabaceae)

The Plant Vigor Hypothesis (PVH) proposes that natural selection on female oviposition choice results from higher fitness of larvae on more vigorous and larger plant modules. For six consecutive years we tested the PVH predictions by investigating the effect of shoot size of Bauhinia brevipes (Fabaceae) on the oviposition preference and offspring survival of the gall-midge Schizomyia macrocapillata (Diptera: Cecidomyiidae). Additionally, we analyzed the effects of bottom-up and top-down mortality forces on the system. The hypersensitive reaction (bottom-up effect) accounted for more than 90% larval mortality of S. macrocapillata, making available few galls to be found and killed by natural enemies (top-down effect). Smaller shoots were always more abundant while longer shoots were rare. Nevertheless, the percent number of galls induced by S. macrocapillata was up to 10-fold greater on the largest shoots, corroborating the preference prediction of the PVH. Schizomyia macrocapillata should use over-exploit larger shoots to maximize the preference for, and consequently increase the performance on these shoots. Our results partially support the performance prediction of the PVH: (1) the observed survival was higher than expected on longer shoots, and (2) the ratio of survival per shoot was positively related with shoot length only in 2 years. Thus, we found a link between female preference and larval performance on large-sized shoots, at least in some years. The gall-midge attack pattern in this study might be an evolved response to maximize the female preference and increased larval performance on longer shoots of the host plant.     

Distribution and frequency of galls induced by Anisodiplosis waltheriae Maia (Diptera: Cecidomyiidae) on the invasive plant Waltheria indica L. (Sterculiaceae)

The frequency of galls induced by Anisodiplosis waltheriae Maia, a recently described species, on Waltheria indica L. was studied. W indica is an invasive weed in regeneration areas of Atlantic Forest in southeastern Brazil. Plants were collected in May 2004 and above-ground biomass, main stem length, number of leaves, number of galls per leaf and leaf area of each individual were recorded. Nearly 90% of all plants and 25% of all leaves were attacked by the gall midge, with an average of 0.67 galls/leaf. Leaf area had a weak effect on gall abundance while the number of leaves had no effect on gall abundance. Only 31% of the variation in gall abundance was explained by plant biomass. Natural enemies killed one third of the sampled galls. Predation accounted for 22.9% of gall mortality, unknown factors killed 7.6%, microhymenopteran parasitoids killed 2.5% and fungi only 1%. Mortality factors were not influenced by leaf area or gall density.     

A review of the biology,ecology and control of saddle gall midge,Haplodiplosis marginata (Diptera: Cecidomyiidae) with a focus on phenological forecasting

Saddle gall midge Haplodiplosis marginata (Diptera: Cecidomyiidae) is a pest of cereals across Europe. The occasional nature of this pest has resulted in limited and sporadic research activity. There remain important gaps in knowledge due either to a genuine lack of research or to previous research being difficult to access. These knowledge gaps make the development of effective control options difficult. Here, we review the existing literature in an attempt to consolidate the information on H. marginata from research which spans several decades and encompasses many different countries. The current distribution and pest status of this insect are updated, along with the methods of cultural and chemical control available to growers. The biology and life history of the insect are described in detail and the ecological processes governing them are discussed. A forecasting model is presented which allows the emergence of this pest in the UK to be predicted from degree day data, and the potential application of this model in management decisions is discussed. Finally, the areas in most need of further research are identified, along with suggestions of how this information can be used to help develop effective and sustainable management solutions for this pest.