Since its accidental introduction from Asia, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has killed millions of ash trees in North America. As it continues to ...spread, it could functionally extirpate ash with devastating economic and ecological impacts. Little was known about EAB when it was first discovered in North America in 2002, but substantial advances in understanding of EAB biology, ecology, and management have occurred since. Ash species indigenous to China are generally resistant to EAB and may eventually provide resistance genes for introgression into North American species. EAB is characterized by stratified dispersal resulting from natural and human-assisted spread, and substantial effort has been devoted to the development of survey methods. Early eradication efforts were abandoned largely because of the difficulty of detecting and delineating infestations. Current management is focused on biological control, insecticide protection of high-value trees, and integrated efforts to slow ash mortality.
Urbanization affects communities of herbivorous arthropods and provides opportunities for dramatic changes in their abundance and richness. Underlying these changes are creation of impervious ...surfaces; variation in the density, diversity, and complexity of vegetation; and maintenance practices including pulsed inputs of fertilizers, water, and pesticides. A rich body of knowledge provides theoretical underpinnings for predicting and understanding impacts of urbanization on arthropods. However, relatively few studies have elucidated mechanisms that explain patterns of insect and mite abundance and diversity across urbanization gradients. Published accounts suggest that responses to urbanization are often taxon specific, highly variable, and linked to properties of urbanization that weaken top-down and/or bottom-up processes, thereby destabilizing populations of herbivores and their natural enemies. In addition to revealing patterns in diversity and abundance of herbivores across urbanization gradients, a primary objective of this review is to examine mechanisms underlying these patterns and to identify potential hypotheses for future testing.
Emerald ash borer (EAB;
Agrilus planipennis
) has killed millions of ash trees and threatens ash throughout North America, and long-term persistence of ash will depend on the potential for ...regeneration. We quantified ash demography, including mortality and regeneration, of
Fraxinus americana
(white ash),
Fraxinus pennsylvanica
(green ash), and
Fraxinus nigra
(black ash) in mixed hardwood forests near the epicenter of the EAB invasion in southeastern Michigan and throughout Ohio. Plots were established across a gradient of ash densities. Ash was the most important species in hydric sites, and ranked second among all species in mesic and xeric sites. In sites nearest the epicenter in Michigan, ash mortality exceeded 99 % by 2009, and few or no newly germinated ash seedlings were observed, leaving only an “orphaned cohort” of established ash seedlings and saplings. As ash mortality increased, the number of viable ash seeds in soil samples decreased sharply, and no viable seeds were collected in 2007 or 2008. In Ohio sites farther from the epicenter, densities of new ash seedlings were much higher in plots with healthy ash trees compared to plots where trees had died. EAB was still present in low densities in Michigan and Ohio stands in 2012 where average mortality of ash was nearly 100 %. The future of ash at these sites will depend on the outcome of the dynamic interaction between the orphaned cohort of previously established ash seedlings and saplings and low density EAB populations.
Alien invasive insects such as gypsy moth, hemlock woolly adelgid, and emerald ash borer continue to disturb the mixed deciduous and hemlock forests of eastern North America by causing wide-scale ...defoliation, decline and/or mortality of their hosts. Some of the most devastating species are spreading in “defense free space”, causing extensive mortality of hosts that are inherently susceptible, perhaps due to their lack of coevolutionary history with the invader. These disturbances have altered the dynamics of canopy gaps, coarse woody debris, biogeochemical cycling, and ecological interactions among organisms in terrestrial and aquatic systems, with consequent effects on forest composition, structure, and function. Populations of indigenous species specialized to particular habitats and/or host trees are most likely to decrease, while some generalist and opportunistic species may increase in invaded forests, including exotic plants as their facilitation by alien insects sparks an “invasional meltdown”. Although poorly documented, alien insects may induce positive feedback effects on ecological processes and interactions. For example, effects of herbivory on foliar chemistry may indirectly alter tri-trophic interactions of indigenous herbivores on their shared hosts, slow rates of terrestrial nutrient cycling, and decrease productivity of aquatic habitats based on allochthonous inputs. Tactics used to eradicate or suppress alien insects in forests such as insecticide applications, biological control, and silvicultural prescriptions can also have ecological impacts. As alien insects continue to establish and spread in forests of eastern North America, their already pervasive effects on ecological interactions and ecosystem processes will continue to magnify.
The insect midgut and fat body represent major tissue interfaces that deal with several important physiological functions including digestion, detoxification and immune response. The emerald ash ...borer (Agrilus planipennis), is an exotic invasive insect pest that has killed millions of ash trees (Fraxinus spp.) primarily in the Midwestern United States and Ontario, Canada. However, despite its high impact status little knowledge exists for A. planipennis at the molecular level.
Newer-generation Roche-454 pyrosequencing was used to obtain 126,185 reads for the midgut and 240,848 reads for the fat body, which were assembled into 25,173 and 37,661 high quality expressed sequence tags (ESTs) for the midgut and the fat body of A. planipennis larvae, respectively. Among these ESTs, 36% of the midgut and 38% of the fat body sequences showed similarity to proteins in the GenBank nr database. A high number of the midgut sequences contained chitin-binding peritrophin (248)and trypsin (98) domains; while the fat body sequences showed high occurrence of cytochrome P450s (85) and protein kinase (123) domains. Further, the midgut transcriptome of A. planipennis revealed putative microbial transcripts encoding for cell-wall degrading enzymes such as polygalacturonases and endoglucanases. A significant number of SNPs (137 in midgut and 347 in fat body) and microsatellite loci (317 in midgut and 571 in fat body) were predicted in the A. planipennis transcripts. An initial assessment of cytochrome P450s belonging to various CYP clades revealed distinct expression patterns at the tissue level.
To our knowledge this study is one of the first to illuminate tissue-specific gene expression in an invasive insect of high ecological and economic consequence. These findings will lay the foundation for future gene expression and functional studies in A. planipennis.
63 I. 64 II. 64 III. 65 IV. 71 V. 72 75 References 75 SUMMARY: We review the literature on host resistance of ash to emerald ash borer (EAB, Agrilus planipennis), an invasive species that causes ...widespread mortality of ash. Manchurian ash (Fraxinus mandshurica), which coevolved with EAB, is more resistant than evolutionarily naïve North American and European congeners. Manchurian ash was less preferred for adult feeding and oviposition than susceptible hosts, more resistant to larval feeding, had higher constitutive concentrations of bark lignans, coumarins, proline, tyramine and defensive proteins, and was characterized by faster oxidation of phenolics. Consistent with EAB being a secondary colonizer of coevolved hosts, drought stress decreased the resistance of Manchurian ash, but had no effect on constitutive bark phenolics, suggesting that they do not contribute to increased susceptibility in response to drought stress. The induced resistance of North American species to EAB in response to the exogenous application of methyl jasmonate was associated with increased bark concentrations of verbascoside, lignin and/or trypsin inhibitors, which decreased larval survival and/or growth in bioassays. This finding suggests that these inherently susceptible species possess latent defenses that are not induced naturally by larval colonization, perhaps because they fail to recognize larval cues or respond quickly enough. Finally, we propose future research directions that would address some critical knowledge gaps.
Eradication is the deliberate elimination of a species from an area. Given that international quarantine measures can never be 100% effective, surveillance for newly arrived populations of nonnative ...species coupled with their eradication represents an important strategy for excluding potentially damaging insect species. Historically, eradication efforts have not always been successful and have sometimes been met with public opposition. But new developments in our understanding of the dynamics of low-density populations, the availability of highly effective treatment tactics, and bioeconomic analyses of eradication strategies offer new opportunities for developing more effective surveillance and eradication programs. A key component that connects these new developments is the harnessing of Allee effects, which naturally promote localized species extinction. Here we review these developments and suggest how research might enhance eradication strategies.
Ash (Fraxinus spp.) is a dominant tree species throughout urban and forested landscapes of North America (NA). The rapid invasion of NA by emerald ash borer (Agrilus planipennis), a wood-boring ...beetle endemic to Eastern Asia, has resulted in the death of millions of ash trees and threatens billions more. Larvae feed primarily on phloem tissue, which girdles and kills the tree. While NA ash species including black (F. nigra), green (F. pennsylvannica) and white (F. americana) are highly susceptible, the Asian species Manchurian ash (F. mandshurica) is resistant to A. planipennis perhaps due to their co-evolutionary history. Little is known about the molecular genetics of ash. Hence, we undertook a functional genomics approach to identify the repertoire of genes expressed in ash phloem.
Using 454 pyrosequencing we obtained 58,673 high quality ash sequences from pooled phloem samples of green, white, black, blue and Manchurian ash. Intriguingly, 45% of the deduced proteins were not significantly similar to any sequences in the GenBank non-redundant database. KEGG analysis of the ash sequences revealed a high occurrence of defense related genes. Expression analysis of early regulators potentially involved in plant defense (i.e. transcription factors, calcium dependent protein kinases and a lipoxygenase 3) revealed higher mRNA levels in resistant ash compared to susceptible ash species. Lastly, we predicted a total of 1,272 single nucleotide polymorphisms and 980 microsatellite loci, among which seven microsatellite loci showed polymorphism between different ash species.
The current transcriptomic data provide an invaluable resource for understanding the genetic make-up of ash phloem, the target tissue of A. planipennis. These data along with future functional studies could lead to the identification/characterization of defense genes involved in resistance of ash to A. planipennis, and in future ash breeding programs for marker development.
In forest ecosystems, disturbances that cause tree mortality create canopy gaps, increase growth of understory vegetation, and alter the abiotic environment. These impacts may have interacting ...effects on populations of ground‐dwelling invertebrates that regulate ecological processes such as decomposition and nutrient cycling. A manipulative experiment was designed to decouple effects of simultaneous disturbances to the forest canopy and ground‐level vegetation to understand their individual and combined impacts on ground‐dwelling invertebrate communities. We quantified invertebrate abundance, richness, diversity, and community composition via pitfall traps in response to a factorial combination of two disturbance treatments: canopy gap formation via girdling and understory vegetation removal. Formation of gaps was the primary driver of changes in invertebrate community structure, increasing activity‐abundance and taxonomic richness, while understory removal had smaller effects. Families of Collembola and Diplopoda, as well as some families of Coleoptera, increased in combined canopy and understory disturbance treatments, whereas Curculionidae and Nitidulidae were more abundant in undisturbed forest. Gaps increased light availability, height and cover of understory vegetation, and soil moisture levels, and decreased depth and cover of leaf litter compared to undisturbed forest. Decoupling of canopy and understory vegetation disturbances revealed gap formation as an important short‐term driver of ground‐dwelling invertebrate community structure and composition. Our findings increase understanding of how ground‐dwelling invertebrate communities respond to disturbance and inform sustainable management of forest ecosystems to foster biodiversity and resilience.
•Canopy gaps decreased richness and diversity, but had minor effects on composition.•Ash coarse woody debris (CWD) had minimal effects on forest floor invertebrates.•An inverse temporal relationship ...in magnitude effects of gaps and CWD is proposed.
Emerald ash borer (EAB; Agrilus planipennis) is an invasive wood-boring beetle that causes rapid ash tree mortality, widespread formation of canopy gaps, and accumulation of coarse woody debris (CWD) in forests. The objective was to quantify the effects of canopy gaps, ash CWD and their interaction caused by EAB-induced ash mortality on the forest floor invertebrate community. A two-year manipulative experiment was conducted in forests at NASA Plum Brook Research Station in Ohio, USA. Canopy gaps caused by ash mortality decreased the activity-abundance of some taxa, such as Opiliones, Carabidae and several families of Collembola, as well as invertebrate richness and diversity, but had minor effects on community composition. CWD had minimal effects on the forest floor invertebrate community overall, but altered the activity-abundance of several invertebrate groups, and marginally increased taxonomic evenness in closed canopy treatments. The results of this study suggest that the effects of ash mortality on forest floor invertebrate communities change over time due to an inverse temporal relationship in magnitude effects of canopy gaps, which diminish over time, and accumulation of CWD, which increases over time. These findings contribute to our understanding of the effects of disturbance on ecosystems and the cascading ecological impacts of emerald ash borer invasions in forests.
PDF
