Host plant associations and geography interact to shape diversification in a specialist insect herbivore

Disentangling the processes underlying geographic and environmental patterns of biodiversity challenges biologists as such patterns emerge from eco‐evolutionary processes confounded by spatial autocorrelation among sample units. The herbivorous insect, Belonocnema treatae (Hymenoptera: Cynipidae), exhibits regional specialization on three plant species whose geographic distributions range from sympatry through allopatry across the southern United States. Using range‐wide sampling spanning the geographic ranges of the three host plants and genotyping‐by‐sequencing of 1,217 individuals, we tested whether this insect herbivore exhibited host plant‐associated genomic differentiation while controlling for spatial autocorrelation among the 58 sample sites. Population genomic structure based on 40,699 SNPs was evaluated using the hierarchical Bayesian model entropy to assign individuals to genetic clusters and estimate admixture proportions. To control for spatial autocorrelation, distance‐based Moran's eigenvector mapping was used to construct regression variables summarizing spatial structure inherent among sample sites. Distance‐based redundancy analysis (dbRDA) incorporating the spatial variables was then applied to partition host plant‐associated differentiation (HAD) from spatial autocorrelation. By combining entropy and dbRDA to analyse SNP data, we unveiled a complex mosaic of highly structured differentiation within and among gall‐former populations finding evidence that geography, HAD and spatial autocorrelation all play significant roles in explaining patterns of genomic differentiation in B. treatae. While dbRDA confirmed host association as a significant predictor of patterns of genomic variation, spatial autocorrelation among sites explained the largest proportion of variation. Our results demonstrate the value of combining dbRDA with hierarchical structural analyses to partition spatial/environmental patterns of genomic variation.     

Identification of the parasitoid community associated with an outbreaking gall wasp,Zapatella davisae,and their relative abundances in New England and Long Island,New York

Gall wasps (Hymenoptera: Cynipidae) are phytophagous insects that often go unnoticed; however, when they are introduced to a new area or released from their natural enemies, they have the capacity to outbreak and cause extensive foliar damage. One such outbreaking pest, Zapatella davisae (Cynipidae: Cynipini), causes significant damage and mortality to black oak, Quercus velutina, in the northeastern United States. In this study, we aimed to identify the parasitoid community associated with Z. davisae, compare differences in percent parasitism of Z. davisae in Cape Cod and Long Island, and determine which parasitoid species contribute most to parasitism in each region. From both locations, we reared parasitoids, identified morphological groups, analyzed percent parasitism rates for each group, and used DNA barcoding to provide species‐level identifications. On Long Island, there was nearly 100% parasitism in 2015 followed by a near total collapse of the population in 2016. In contrast, parasitism rates were lower and remained consistent on Cape Cod between 2015 and 2016, which may explain the greater canopy damage observed in that region. Species of Sycophila were the dominant parasitoids, with one species Sycophila nr. novascotiae representing ~65% of reared parasitoids from Long Island, and two species of Sycophila (S. nr. novascotiae and S. foliatae) with near equal representations on Cape Cod. In order to manage an insect pest, it is important to understand factors that influence its mortality and survival. An understanding of how these infestations progress overtime can help predict the impact that newer infestations in Nantucket, MA, and coastal Rhode Island will have on black oak populations and will aid in the management of this rapidly spreading gall wasp pest.     

Stem galls affect oak foliage with potential consequences for herbivory

Abstract.   1. On two dates, foliar characteristics of pin oak, Quercus palustris , infested with stem galls caused by the horned oak gall, Callirhytis cornigera , were investigated, and the consequences for subsequent herbivory assessed.
2. Second-instar caterpillars of the gypsy moth, Lymantria dispar , preferred foliage from ungalled trees.
3. Ungalled trees broke bud earlier than their galled counterparts.
4. Galled trees produced denser leaves with higher nitrogen and tannin concentrations, but foliar carbohydrates did not differ among galled and ungalled trees.
5. Concentrations of foliar carbohydrates in both galled and ungalled trees increased uniformly between the two assay dates. Nitrogen concentrations were greater in leaves from galled trees, and decreased uniformly in galled and ungalled trees over time. Foliar tannins were also greater in foliage from galled trees early in the season; however, foliar tannins declined seasonally in galled tissue so that by the second assay date there was no difference in tannin concentrations between galled and ungalled foliage.
6. In spite of differences in foliar characteristics, performance of older, fourth instar gypsy moth caterpillars did not differ between galled and ungalled trees.     

Evolutionary shifts between host oak sections and host-plant organs in Andricus gallwasps

Abstract.— Gall-inducing insects have especially intimate interactions with their host plants and generally show great specificity with regard to both the host-plant species and the organ (e.g. flower, leaf) galled. However, the relative roles of shifts between host species and between host-plant organs in the diversification of gall-inducers are uncertain. We employ a novel and general maximum-likelihood approach to show that shifts between host-plant organs occur at a significantly greater rate than shifts between host oak sections in European Andricus gallwasps. This suggests that speciation has more often been associated with gall location shifts than with colonization of new host-plant species, and implies that it may be easier for gall-inducers to colonize new plant organs than new plant species.
Andricus gallwasps have complex life cycles, with obligate alternation of sexual and parthenogenetic generations. Our phylogenetic analyses show that a life cycle with both generations galling white oaks (section Quercus ) is ancestral, with a single shift of the sexual generation onto black oaks (section Cerris ) to generate a clade with a novel host-alternating life cycle. This new life cycle provided the opportunity for further speciation, but may have also increased the risk of extinction of one or both generations by the demographic requirement for co-existence of both host-plant groups. In summary, it appears that Andricus gallwasp radiation may be a two-level process. Speciation events often involve shifts in gall location on the same host species. However, there are only so many ways to gall an oak, and rare shifts to new oak sections may contribute greatly to long-term diversification by opening up whole new adaptive zones.     

Impact of prescribed burning on endophytic insect communities of prairie perennials (Asteraceae: Silphium spp.)

Prescribed burning currently is used to preserve endemicity of plant communities in remnant tallgrass prairies. Although some types of arthropods benefit from changes in plant communities brought about by burning, other species that are endemic to prairies may be threatened. Because they inhabit the fuel layer of prairies, endophytic insects would seem particularly susceptible to this management tactic. In this paper, we assess the impact of prescribed burning on endophytic insect communities inhabiting stems of Silphium laciniatum L. and S. terebinthinaceum Jacquin (Asteraceae), endemic prairie plants. Populations of these insects were decimated by burning, with mortality approaching 100% in most cases. Their populations nevertheless began to rebound within a single growing season, with densities moderately but significantly reduced 1 year after the burn. Even when a prairie remnant was completely incinerated, plant stems were recolonized by insects within one growing season. Our findings suggest that sufficient numbers of endophytic insects survive burns in remains of Silphium to recolonize burned areas the following year.     

Invasion by the chestnut gall wasp in Italy causes significant yield loss in Castanea sativa nut production

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Trade‐off between fecundity and survival generates stabilizing selection on gall size

Complex interactions within multitrophic communities are fundamental to the evolution of individual species that reside within them. One common outcome of species interactions are fitness trade‐offs, where traits adaptive in some circumstances are maladaptive in others. Here, we identify a fitness trade‐off between fecundity and survival in the cynipid wasp Callirhytis quercusbatatoides that induces multichambered galls on the stem of its host plant Quercus virginiana. We first quantified this trade‐off in natural populations by documenting two relationships: a positive association between the trait gall size and fecundity, as larger galls contain more offspring, and a negative association between gall size and survival, as larger galls are attacked by birds at a higher rate. Next, we performed a field‐based experimental evolution study where birds were excluded from the entire canopy of 11 large host trees for five years. As a result of the five‐year release from avian predators, we observed a significant shift to larger galls per tree. Overall, our study demonstrates how two opposing forces of selection can generate stabilizing selection on a critical phenotypic trait in wild populations, and how traits can evolve rapidly in the predicted direction when conditions change.     

Environmental thermal levels affect the phenological relationships between the chestnut gall wasp and its parasitoids

Studies of thermal level‐related asynchrony in a host–parasitoid relationship are necessary to understand the effects of climate change on new host–parasitoid interactions. In the Asian chestnut gall wasp Dryocosmus kuriphilus (Hymenoptera: Cynipidae) and its Chalcidoidea parasitoids, phenological synchrony is assumed to be weather‐dependent in a new area of expansion. To evaluate the effects of environmental thermal regimes on the host, a phenology model for different cynipid stages (larvae, pupae, adults, and adult emergence) and a host–parasitoid phenological estimator are developed in three chestnut fields during two successive growth seasons and subsequently validated in areas with chestnut fields at two different altitudes. Comparisons of the timings of the juvenile and adult stages with those of the parasitoid complex demonstrate that the shortest period of occurrence for cynipids within galls has negative effects on the host–parasitoid relationships at higher temperature levels, thereby increasing phenological asynchrony for some parasitoids species. Reducing the development time of pupae and adults decreases the likelihood of success for some parasitoid species at higher temperature levels. We also record the extension of the gall wasp development time (approximately 15 days) at higher altitudes (linked to a lower mean temperature of approximately 1.5 °C). These results highlight how parasitization on the new hosts is dependent on the host phenology and, in the present study, is limited by the short duration of the presence of the host in galls, which could explain the considerable differences in cynipid gall wasp parasitization recorded at different altimeters.     

Current state of knowledge of the Korean Cynipini: subspecies description,new combinations and checklist (Hymenoptera,Cynipidae)

Prior to this study, only 8 species of oak gall-forming wasps were known from Korea. After material examination and an exhaustive bibliographic revision, a total of 17 oak gall wasps species have been reported. The following species correspond to new records: Andricus hakonensis (Ahmead); A. pseudocurvator Tang & Melika; A. songshui Tang & Melika; Cerroneuroterus folimargo (Monzen); C. monzeni (Dettmer); C. vonkuenburgi (Dettmer); Neuroterus hakonensis Ashmead, Trichagalma acutissimae (Monzen), T. formosana Melika & Tang. It was impossible to obtain adult specimens for all the species, and and the following species are recorded only based on galls: Andricus kashiwaphilus Abe; A. mukaigawae (Mukaigawa); A. pseudoflos (Monzen); A. targionii Kieffer; Cerroneuroterus folimargo (Monzen); C. monzeni (Dettmer); Neuroterus hakonensis Ashmead; Trichagalma acutissimae (Monzen). The species Dryocosmus kunugiphagus Ide & Abe, 2015 is transferred to the genus Plagiotrochus: Plagiotrochus kunugiphagus (Ide & Abe, 2015) n. comb. Additionally a new subspecies of Trichagalma formosana is described to include the material from Korea and Mainland China, closely related but morphologically different to the Taiwan populations. For each species live cycle, morphology of galls, hosts, phenology, and distribution are commented. In some of them taxonomic or morphological comments are mentioned and a key of agamic Trichagalma with dark smoky pigmented spots in the forewing is added. All galls are illustrated. An up-to-date checklist of the Korean Cynipini is provided. The previously reported species from Korea (Andricus noliquercicola Shinji, 1938; Aphelonyx crispulae Mukaigawae, 1920; Neuroterus narae Shinji, 1941 and Neuroterus nawai Ashmead, 1904) are here excluded from the Korean species list.