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      α-Copaene is an attractant, synergistic with quercivorol, for improved detection of Euwallacea nr. fornicatus (Coleoptera: Curculionidae: Scolytinae)

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          Abstract

          The tea shot-hole borer, Euwallacea fornicatus Eichhoff, is an ambrosia beetle endemic to Asia and a pest of commercial tea, Camellia sinensis (L.) Kuntze. Recently, a complex of species morphologically similar to E. fornicatus has been recognized, which includes new pests established in Israel and the USA, both in California and Florida. Collectively termed E. nr. fornicatus, these cryptic species carry symbiotic Fusarium spp. fungi, some of which cause dieback disease in susceptible hosts, which include avocado, Persea americana Miller. Due to the threat to this economically important crop, research was initiated to evaluate efficacy of kairomone-based lures for detection of the beetle in Florida (termed the Florida tea shot hole borer, FL-TSHB). A series of field tests were conducted in 2016 in commercial avocado groves known to have FL-TSHB at various population levels. All tests evaluated lures containing quercivorol ( p-menth-2-en-1-ol) and α-copaene, presented separately and in combination; and one test evaluated effect of trap type on beetle captures. In addition, electroantennography (EAG) was used to quantify female olfactory responses to lure emissions. This study identified (-)-α-copaene as a new attractant for FL-TSHB, equivalent in efficacy to quercivorol (the standard lure for Euwallacea detection in the USA); however, the combination of lures captured significantly more FL-TSHB than either lure alone. This combination resulted in synergistic attraction at two field sites and additive attraction at a third site. Sticky panel traps captured more FL-TSHB than comparably-baited Lindgren funnel traps. Females engaged in host-seeking flight from 11:00 to 16:00 hr (EST), with peak numbers observed between 12:00 and 13:00 hr. EAG analyses confirmed olfactory chemoreception of both kairomones, with a higher response elicited with the combination of volatiles. Results indicate that detection of pest E. nr. fornicatus in Florida can be improved by using a two-component lure consisting of p-menth-2-en-1-ol and (-)-α-copaene.

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          Review of American Xyleborina (Coleoptera: Curculionidae: Scolytinae) Occurring North of Mexico, with an Illustrated Key

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            The sudden emergence of pathogenicity in insect-fungus symbioses threatens naive forest ecosystems.

            Invasive symbioses between wood-boring insects and fungi are emerging as a new and currently uncontrollable threat to forest ecosystems, as well as fruit and timber industries throughout the world. The bark and ambrosia beetles (Curculionidae: Scolytinae and Platypodinae) constitute the large majority of these pests, and are accompanied by a diverse community of fungal symbionts. Increasingly, some invasive symbioses are shifting from non-pathogenic saprotrophy in native ranges to a prolific tree-killing in invaded ranges, and are causing significant damage. In this paper, we review the current understanding of invasive insect-fungus symbioses. We then ask why some symbioses that evolved as non-pathogenic saprotrophs, turn into major tree-killers in non-native regions. We argue that a purely pathology-centred view of the guild is not sufficient for explaining the lethal encounters between exotic symbionts and naive trees. Instead, we propose several testable hypotheses that, if correct, lead to the conclusion that the sudden emergence of pathogenicity is a new evolutionary phenomenon with global biogeographical dynamics. To date, evidence suggests that virulence of the symbioses in invaded ranges is often triggered when several factors coincide: (i) invasion into territories with naive trees, (ii) the ability of the fungus to either overcome resistance of the naive host or trigger a suicidal over-reaction, and (iii) an 'olfactory mismatch' in the insect whereby a subset of live trees is perceived as dead and suitable for colonization. We suggest that individual cases of tree mortality caused by invasive insect-fungus symbionts should no longer be studied separately, but in a global, biogeographically and phylogenetically explicit comparative framework.
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              Fusarium euwallaceae sp. nov.--a symbiotic fungus of Euwallacea sp., an invasive ambrosia beetle in Israel and California.

              The invasive Asian ambrosia beetle Euwallacea sp. (Coleoptera, Scolytinae, Xyleborini) and a novel Fusarium sp. that it farms in its galleries as a source of nutrition causes serious damage to more than 20 species of live trees and pose a serious threat to avocado production (Persea americana) in Israel and California. Adult female beetles are equipped with mandibular mycangia in which its fungal symbiont is transported within and from the natal galleries. Damage caused to the xylem is associated with disease symptoms that include sugar or gum exudates, dieback, wilt and ultimately host tree mortality. In 2012 the beetle was recorded on more than 200 and 20 different urban landscape species in southern California and Israel respectively. Euwallacea sp. and its symbiont are closely related to the tea shot-hole borer (E. fornicatus) and its obligate symbiont, F. ambrosium occurring in Sri Lanka and India. To distinguish these beetles, hereafter the unnamed xyleborine in Israel and California will be referred to as Euwallacea sp. IS/CA. Both fusaria exhibit distinctive ecologies and produce clavate macroconidia, which we think might represent an adaption to the species-specific beetle partner. Both fusaria comprise a genealogically exclusive lineage within Clade 3 of the Fusarium solani species complex (FSSC) that can be differentiated with arbitrarily primed PCR. Currently these fusaria can be distinguished only phenotypically by the abundant production of blue to brownish macroconidia in the symbiont of Euwallacea sp. IS/CA and their rarity or absence in F. ambrosium. We speculate that obligate symbiosis of Euwallacea and Fusarium, might have driven ecological speciation in these mutualists. Thus, the purpose of this paper is to describe and illustrate the novel, economically destructive avocado pathogen as Fusarium euwallaceae sp. nov. S. Freeman et al.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                13 June 2017
                2017
                : 12
                : 6
                : e0179416
                Affiliations
                [1 ]United States Department of Agriculture, Agricultural Research Service, Subtropical Horticulture Research Station, Miami, FL, United States of America
                [2 ]United States Department of Agriculture, Agricultural Research Service, Beltsville Area Research Center, Electron and Confocal Microscopy Unit, Beltsville, MD, United States of America
                [3 ]University of Florida, Tropical Research and Education Center, Homestead, FL, United States of America
                Universidade Federal de Vicosa, BRAZIL
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                • Conceptualization: PK WM DC.

                • Data curation: PK TN.

                • Formal analysis: PK.

                • Investigation: PK DO WM TN GB ES NT.

                • Methodology: PK WM GB NT.

                • Project administration: PK.

                • Resources: PK DO GB NT DC.

                • Supervision: PK.

                • Visualization: PK GB.

                • Writing – original draft: PK.

                • Writing – review & editing: PK DO NT DC.

                Author information
                http://orcid.org/0000-0003-4425-0733
                Article
                PONE-D-17-15817
                10.1371/journal.pone.0179416
                5469513
                28609448
                4e41f676-c54a-40f0-9c26-5b0edabbe04a

                This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 24 April 2017
                : 30 May 2017
                Page count
                Figures: 6, Tables: 2, Pages: 20
                Funding
                This research was supported by the United States Department of Agriculture, Agricultural Research Service (USDA-ARS) Appropriated Funds (Mitigation of the Invasive Pest Threat from the American Tropics and Subtropics) and by an appointment to the ARS Research Participation agreement between the U.S. Department of Energy (DOE) and the U.S. Department of Agriculture (USDA). ORISE is managed by ORAU under DOE contract number DE-SC0014664. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Biology and Life Sciences
                Organisms
                Animals
                Invertebrates
                Arthropoda
                Insects
                Beetles
                Research and Analysis Methods
                Research Design
                Field Tests
                People and places
                Geographical locations
                North America
                United States
                Florida
                Biology and Life Sciences
                Physiology
                Biological Locomotion
                Flight (Biology)
                Insect Flight
                Medicine and Health Sciences
                Physiology
                Biological Locomotion
                Flight (Biology)
                Insect Flight
                Biology and Life Sciences
                Physiology
                Biological Locomotion
                Flight (Biology)
                Medicine and Health Sciences
                Physiology
                Biological Locomotion
                Flight (Biology)
                Biology and Life Sciences
                Organisms
                Plants
                Trees
                Biology and Life Sciences
                Zoology
                Animal Anatomy
                Antennae (Animal Physiology)
                Biology and Life Sciences
                Zoology
                Animal Physiology
                Antennae (Animal Physiology)
                Biology and Life Sciences
                Microbiology
                Medical Microbiology
                Microbial Pathogens
                Fungal Pathogens
                Fusarium
                Medicine and Health Sciences
                Pathology and Laboratory Medicine
                Pathogens
                Microbial Pathogens
                Fungal Pathogens
                Fusarium
                Biology and Life Sciences
                Mycology
                Fungal Pathogens
                Fusarium
                Custom metadata
                All relevant data are within the paper and its Supporting Information files.

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