Structure and development of ‘witches' broom’ galls in reproductive organs of Byrsonima sericea (Malpighiaceae) and their effects on host plants

Galls are anomalies in plant development of parasitic origin that affect the cellular differentiation or growth and represent a remarkable plant–parasite interaction. Byrsonima sericea DC. (Malpighiaceae) is a super host of several different types of gall in both vegetative and reproductive organs. 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: the ‘witches' broom’ galls developed in floral structures of B. sericea. The unaffected inflorescences are characterised by a single indeterminate main axis with spirally arranged flower buds. The flower buds developed five unaffected brownish hairy sepals and five pairs of elliptical yellow elaiophores, five yellow fringed petals, 10 stamens and a pistil with superior tricarpellar and trilocular ovary. The affected inflorescences showed changes in architecture, with branches arising from the main axis and flower buds. The flower buds exhibited several morphological and anatomical changes. The sepals, petals and carpels converted into leaf‐like structures after differentiation. Stamens exhibited degeneration of the sporogenous tissue and structures containing hyphae and spores. The gynoecium did not develop, forming a central meristematic region, from which emerges the new inflorescence. In this work, we discuss the several changes in development of reproductive structures caused by witches' broom galls and their effects on reproductive success of the host plants.     

Leaf gall polymorphism and molecular phylogeny of a new Bruggmanniella species (Diptera: Cecidomyiidae: Asphondyliini) associated with Litsea acuminata (Lauraceae) in Taiwan,with ecological comparisons and a species description

Two types of cecidomyiid leaf galls, cup‐shaped and umbrella‐shaped, occur on Litsea acuminata (Lauraceae) in Taiwan. Based on the concept of gall shapes as “extended phenotypes” of gall inducers, these two types could be induced by different gall midge species. However, galls with intermediate shapes between the two types were recently discovered, which implies that possible genetic exchanges occur between the gall inducers of both types. To clarify the taxonomic status of gall midges responsible for the two types of galls on L. acuminata, we undertook taxonomic, molecular phylogenetic and ecological studies. Our findings show that the two gall types are induced by the same Bruggmanniella species and the species is new to science. We describe the species forming this range of galls as Bruggmanniella litseae sp. n. , and compare their geographical distribution, galling position and morphometry. Based on our results, a possible evolutionary scenario of B. litseae sp. n. is discussed.     

New gall midge taxa of the subfamilies porricondylinae and lasiopterinae (Diptera,Cecidomyiidae) from the Rovno amber

Five new gall midge taxa of the subfamilies Porricondylinae and Lasiopterinae are described from the Late Eocene Rovno amber: Adsumyia integra gen. et sp. nov. (Dicerurini), Gratomyia inexigentis gen. et sp. nov. (Holoneurini), Winnertzia recusata sp. nov. (Winnertziini), Kovaleviola injusta gen. et sp. nov., and Spungisiola insuperabilis gen. et sp. nov. (Brachineurini).     

In vitro allelopathic effects of extracts and amenthoflavone from Byrsonima crassa (Malpighiaceae)

Abstract

Extracts and pure amenthoflavone isolated from Byrsonima crassa (Malpighiaceae), a shrub growing in the semi-arid region of Brazil Cerrado, were evaluated in vitro, at different doses, for their effects on tomato seed germination and subsequent growth of seedlings. A hydromethanolic extract showed general stimulatory effects. The EtOAc extract stimulated root elongation and root weight of tomato; shoot elongation was inhibited, while shoot weight was not altered. The pure amenthoflavone isolated from the plant, stimulated shoot elongation at concentrations ranging between 10^?4 M and 10^?6 M.     

Morphological,genetic and symptomatic identification of an invasive jujube pest in Korea,Dasineura jujubifolia Jiao & Bu (Diptera: Cecidomyiidae)

Correct identification of invasive species is an important step for further management of this species. New type of leafcurling gall midge feeding on jujube, Ziziphus jujube Miller (Rhamnaceae), was discovered in Korea in 2011. At that time, this species was not identified as specific level, so it was recorded as Dasineura sp. Since then, the jujube gall midge has become a serious alien insect pest in Korea. In this study, this jujube gall midge was surveyed by collecting damaged leaf samples from different major jujube-producing regions in Korea. Morphological, genetic and symptomatic identification of this invasive jujube pest identified as Dasineura jujubifolia Jiao & Bu. Here we provided its taxonomic status, geographical distribution, morphological characteristics of all stages and infestation symptoms. In addition to classical morphological characters, mitochondrial COI barcoding sequences were generated for several specimens. The possible pathway of invasion and subsequent socioeconomic consequences were discussed.     

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.     

Detection of two new cryptic species of Kiefferia (Diptera: Cecidomyiidae) by means of morphological,molecular and ecological studies

Morphological, molecular and ecological studies revealed that Kiefferia Mik, 1985 (Diptera: Cecidomyiidae) is not a monotypic genus. Two new species, Kiefferia ezoensis sp. nov. and K. olla sp. nov. from Japan and Korea are added to the known European species, Kiefferia pericarpiicola. The new and known species are distinguishable from each other mainly by the shape of the larval sternal spatula. Molecular data support the results of morphological classification. Species of Kiefferia induce fruit galls on various plant species of Apiaceae. We found that K. ezoensis and K. olla utilized exclusively Angelica ursina and Oenanthe javanica, respectively. In contrast, we listed 37 apiaceous species belonging to 23 genera as hosts for K. pericarpiicola from previously published reports, suggesting the existence of additional undescribed species. Life history patterns of the three species are distinctly different from each other. Mature larvae of K. ezoensis and K. olla drop to the ground with the fruit gall in October and September, respectively, whereas mature larvae of K. pericarpiicola quit the galls and drop to the ground in August. A key to the Kiefferia species is provided based on morphological features and information on life history patterns and host ranges.     

Description,phenology and biology of Zelostemma chionochloae Buhl sp. nov., a platygastrid parasitoid of Eucalyptodiplosis chionochloae (Diptera: Cecidomyiidae) in New Zealand

Abstract

A platygastrid wasp that feeds on the snow tussock flower midge in the inflorescences of New Zealand snow tussock grasses (Chionochloa spp.) is formally described. Zelostemma chionochloae is a specialist natural enemy of Eucalyptodiplosis chionochloae Kolesik which is the most ubiquitous and sophisticated seed predator of Chionochloa. Z. chionochloae is a koinobiont parasitoid and some larvae enter prolonged diapause inside their host for at least 2 years. Methods for adult rearing are described. The phenology of Z. chionochloae is highly synchronised with its host even after 2 years in diapause. Parasitism levels were found to differ between years and elevations, while sex ratios differed among years. Z chionochloae probably suffers inter‐specific competition with another host‐specific hymenopteran parasitoid (Gastrancistrus sp.) which also parasitises E. chionochloae.     

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.     

Influence of synchronization between adult emergence and host plant phenology on the population density of Pseudasphondylia neolitseae (Diptera: Cecidomyiidae) inducing leaf galls on Neolitsea sericea (Lauraceae)

Synchronization between the appearance of herbivorous insects and their host-plant phenology is a critical event, especially for short-lived insects such as gall midges. We studied a natural population of Pseudasphondylia neolitseae (Diptera: Cecidomyiidae) that induces leaf galls on Neolitsea sericea (Lauraceae) to evaluate the effect of synchronization on gall density in the subsequent generation. To do so, we combined quantitative data on host resources with time lag between emergence and host-available seasons. The gamma distribution model was applied to the emergence curve of P. neolitseae and the normal distribution model to the daily changes in the number of host buds suitable for oviposition; the latter model was transformed into an available-resource curve based on the mean number of host buds required for a single female to realize her eggs. By superimposing the emergence curve on the available-resource curve and calculating overlapped area, the degree of synchronization was evaluated more accurately than previous studies, which had treated only the time lag. The number of females that synchronized with host buds affected gall density in the next generation.     

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.     

Quantitative differences detected in the histology of galls induced by the same aphid species in different varieties of the same host

• Plant galls are abnormal growths caused by an inducer that determines their morphology and anatomy. We qualitatively and quantitatively compared the histological anatomy of five aphid species (Paracletus cimiciformis, Forda marginata, Forda formicaria, Baizongia pistaciae and Geoica wertheimae) that induce galls in Pistacia terebinthus shrubs growing in Israel. We also quantitatively compared these galls to those that the aphids create on the same host in Spain.
• Histological study was conducted following methods described previously by the authors.
• Quantitative differences among the galls were found in five of 12 common anatomical traits: gall thickness, stomatal number in the epidermis‐air, size of vascular bundles, distance of phloem ducts from the lumen and number of intraphloematic schizogenous ducts. Other structures were particular to one or some species: number of cracks in the epidermis–lumen, a sclereid layer, trichomes and microcrystal inclusions. Fisher's tests of combined probabilities showed that the galls induced in Israel were statistically different from those in Spain. In particular, the number of intraphloematic schizogenous ducts was higher in the galls induced in P. terebinthus in Israel. Such differences were also found in other traits related to defence of the gall inhabitant.
• In conclusion, while the gall shape and size are determined mainly by the cecidogenic insect, it seems that the host plant also plays an important role in determining the number/size of quantitative traits, in this case mainly protective structures.

     

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.     

Pathogen effects on vegetative and floral odours mediate vector attraction and host exposure in a complex pathosystem

Pathogens can alter host phenotypes in ways that influence interactions between hosts and other organisms, including insect disease vectors. Such effects have implications for pathogen transmission, as well as host exposure to secondary pathogens, but are not well studied in natural systems, particularly for plant pathogens. Here, we report that the beetle‐transmitted bacterial pathogen Erwinia tracheiphila – which causes a fatal wilt disease – alters the foliar and floral volatile emissions of its host (wild gourd, Cucurbita pepo ssp. texana) in ways that enhance both vector recruitment to infected plants and subsequent dispersal to healthy plants. Moreover, infection by Zucchini yellow mosaic virus (ZYMV), which also occurs at our study sites, reduces floral volatile emissions in a manner that discourages beetle recruitment and therefore likely reduces the exposure of virus‐infected plants to the lethal bacterial pathogen – a finding consistent with our previous observation of dramatically reduced wilt disease incidence in ZYMV‐infected plants.     

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     

Response of two parasitoid species (Hymenoptera: Braconidae,Figitidae) to tephritid host and host food substrate cues

The Neotropical‐native figitid Aganaspis pelleranoi (Brèthes) and the Asian braconid Diachasmimorpha longicaudata (Ashmead) are two parasitoids of Tephritidae fruit flies with long and recent, respectively, evolutionary histories in the Neotropics. Both species experienced a recent range of overlap. In Argentina, A. pelleranoi is a potential species in biological control programs against the pestiferous tephritid species, Anastrepha fraterculus (Wiedemann) and Ceratitis capitata (Wiedemann), whereas D. longicaudata is already used in open‐field releases against Medfly in central‐western Argentina. To characterize the host‐foraging strategies of A. pelleranoi and D. longicaudata, olfactometer experiments were conducted comparing responses to C. capitata and A. fraterculus larvae, in two kinds of food substrate: fruit and artificial larval medium. To control the possible influence of host larvae used for parasitoid rearing on olfactory response, two strains of both parasitoid species, reared on both tephrtid species, were studied. Volatiles directly emanating either from A. fraterculus or C. capitata larvae may be detected by both A. pelleranoi and D. longicaudata, although chemical stimuli originating from the combination of host larvae and the habitat of the host were preferred. However, olfactory cues associated with host larvae probably play a relevant role in host searching behaviour of A. pelleranoi, whereas for D. longicaudata, the host‐habitat olfactory stimuli would be highly essential in short‐range host location. The strain of the parasitoids did not affect host search ability on the two tephritid species evaluated. These evidences are relevant for mass production of both parasitoids and their impact following open‐field augmentative releases.     

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).