Pine-feeding bark beetles (Coleoptera: Scolytidae) interact chemically with their host pines (Coniferales: Pinaceae) via the behavioral, physiological, and biochemical effects of one class of isoprenoids, the monoterpenes and their derivatives. Pine monoterpenes occur in the oleoresin and function as behaviorally active kairomones for pine bark beetles and their predators, presenting a classic example of tritrophic chemical communication. The monoterpenes are also essential co-attractants for pine bark beetle aggregation pheromones. Ironically, pine monoterpenes are also toxic physiologically to bark beetles at high vapor concentrations and are considered an important component of the defense of pines. Research over the last 30 years has demonstrated that some bark beetle aggregation pheromones arise through oxygenation of monoterpenes, linking pheromone biosynthesis to the host pines. Over the last 10 years, however, several frequently occurring oxygenated monoterpene pheromone components (e.g., ipsenol, ipsdienol and frontalin) have also been shown to arise through highly regulated de novo pathways in the beetles (reviewed in Seybold and Tittiger, 2003). The most interesting nexus between these insects and their plant hosts involves the latestage reactions in the monoterpenoid biosynthetic pathway, during which isomeric dimethylallyl diphosphate and isopentenyl diphosphate are ultimately elaborated to stereospecific monoterpenes in the trees and to hydroxylated monoterpenes or bicyclic acetals in the insects. There is signal stereospecificity in both production of and response to the monoterpenoid aggregation pheromones of bark beetles and in response to the monoterpenes of the pines. In the California fivespined ips, Ips paraconfusus, we Dedicated to Professor David L. Wood on the occasion of his 75th birthdayÓ Springer Science+Business Media B.V. 2006have discovered a number of cytochome P450 genes that have expression patterns indicating that they may be involved in detoxifying monoterpene secondary metabolites and/or biosynthesizing pheromone components. Both processes result in the production of oxygenated monoterpenes, likely with varying degrees of stereospecificity. A behavioral analysis of the stereospecific response of I. paraconfusus to its pheromone is providing new insights into the development of an efficacious bait for the detection of this polyphagous insect in areas outside the western United States. In contrast, a Eurasian species that has arrived in California, the Mediterranean pine engraver, Orthotomicus (Ips) erosus, utilizes both a monoterpenoid (ipsdienol) and a hemiterpenoid (2-methyl-3-buten-2-ol) in its pheromone blend. The stereospecificity of the response of O. erosus to the monoterpenoid appears to be the key factor to the improved potency of the attractant bait for this invasive species.

Ponderosa pine and dry mixed-conifer forests in the Southwest United States are experiencing, or have become increasingly susceptible to, large-scale severe wildfire, insect, and disease episodes resulting in altered plant and animal demographics, reduced productivity and biodiversity, and impaired ecosystem processes and functions. We present a management framework based on a synthesis of science on forest ecology and management, reference conditions, and lessons learned during implementations of our restoration framework. Our framework focuses on the restoration of key elements similar to the historical composition and structure of vegetation in these forests: (1) species composition;(2) groups of trees; (3) scattered individual trees; (4) grass-forb-shrub interspaces; (5) snags, logs, and woody debris; and (6) variation in the arrangements of these elements in space and time. Our framework informs management strategies that can improve the resiliency of frequent-fire forests and facilitate the resumption of characteristic ecosystem processes and functions by restoring the composition, structure, and spatial patterns of vegetation. We believe restoration of key compositional and structural elements on a per-site basis will restore resiliency of frequent-fire forests in the Southwest, and thereby position them to better resist, and adapt to, future disturbances and climates.

Utley, C, Nguyen, T., Roubtsova, T, Coggeshall, M., Ford, T. N. 2013. Susceptibility of walnut and hickory species to Geosmithia mórbida. Plant Dis. 97:601-607.Thousand cankers disease (TCD) of walnut is a result of feeding in the phloem by the walnut twig beetle (WTB), Pityophthorus juglandis, and subsequent canker formation caused by Geosmithia mórbida around galleries. TCD has caused extensive morbidity and mortality to Juglans nigra in the western United States and, in 2010, was discovered in the eastern United States, where the tree is a highly valuable timber resource. WTB and G. mórbida also have been found in / regia orchards throughout major production areas in California, and the numbers of damaged trees are increasing. We tested the susceptibility of walnut and hickory species to G. mórbida in greenhouse and field studies. Carya illinoinensis, C. aquatica, and C. ovata were immune. All wal-nut species tested, including / ailantifolia, J. californica, J. cinérea, J. hindsii, J. major, J. mandshurica, J. microcarpa, J. nigra, and J. regia, developed cankers following inoculation with G. mórbida. J. nigra was the most susceptible, whereas / major, a native host of the WTB and, presumably, G. mórbida, had smaller and more superficial cankers. Canker formation differed among maternal half-sibling families of / nigra and / cinérea, indicating genetic variability in resistance to G. mórbida. Our inoculation studies with G. mórbida have corroborated many of the field observations on susceptibility of walnut and hickory species to TCD, although the ability of the WTB to successfully attack and breed in walnut is also an important component in TCD resistance.

The ascomycete Geosmithia morbida and the walnut twig beetle Pityophthorus juglandis are associated with thousand cankers disease of Juglans (walnut) and Pterocarya (wingnut). The disease was first reported in the western United States (USA) on several Juglans species, but has been found more recently in the eastern USA in the native range of the highly susceptible Juglans nigra. We performed a comprehensive population genetic study of 209 G. morbida isolates collected from Juglans and Pterocarya from 17 geographic regions distributed across 12 U.S. states. The study was based on sequence typing of 27 single nucleotide polymorphisms from three genomic regions and genotyping with ten microsatellite primer pairs. Using multilocus sequence-typing data, 197 G. morbida isolates were placed into one of 57 haplotypes. In some instances, multiple haplotypes were recovered from isolates collected on the same tree. Twenty-four of the haplotypes (42%) were recovered from more than one isolate; the two most frequently occurring haplotypes (H02 and H03) represented 36% of all isolates. These two haplotypes were abundant in California, but were not recovered from Arizona or New Mexico. G. morbida population structure was best explained by four genetically distinct groups that clustered into three geographic regions. Most of the haplotypes isolated from the native range of J. major (Arizona and New Mexico) were found in those states only or present in distinct genetic clusters. There was no evidence of sexual reproduction or genetic recombination in any population. The scattered distribution of the genetic clusters indicated that G. morbida was likely disseminated to different regions at several times and from several sources. The large number of haplotypes observed and the genetic complexity of G. morbida indicate that it evolved in association with at least one Juglans spp. and the walnut twig beetle long before the first reports of the disease.

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