The ambrosia beetle–fungus farming symbiosis is more heterogeneous than previously thought. There is not one but many ambrosia symbioses. Beetle-fungus specificity is clade dependent and ranges from strict to promiscuous. Each new origin has evolved a new mycangium. The most common relationship with host trees is colonization of freshly dead tissues, but there are also parasites of living trees, vectors of pathogenic fungi, and beetles living in rotten trees with a wood-decay symbiont. Most of these strategies are driven by fungal metabolism whereas beetle ecology is evolutionarily more flexible. The ambrosia lifestyle facilitated a radiation of social strategies, from fungus thieves to eusocial species to communities assembled by attraction to fungal scent. Although over 95% of the symbiotic pairs are economically harmless, there are also three types of pest damage: tree pathogen inoculation, mass accumulation on susceptible hosts, and structural damage. Beetles able to colonize live tree tissues are most likely to become invasive pests.
Bark and ambrosia beetles (Scolytinae) are the most successful group of invasive wood borers worldwide, and the most invasive among them are species in the tribe Xyleborini. This haplodiploid, highly inbred, fungus-farming group is represented by 30 non-native species in North America, of which at least five are serious pests. The few identification resources for Xyleborini that exist are becoming outdated due to new species arrivals and nomenclatural changes. Here we present a new comprehensive key to Xyleborini currently known from the continental United States. Compared to the previous key, the following species have been added to the North American fauna:Ambrosiodmusminor(Stebbing),Ambrosiophilusnodulosus(Eggers),AnisandrusmaicheKurentsov,Coptoboruspseudotenuis(Schedl),Cyclorhipidionfukiense(Eggers),DryocoetoidesreticulatusAtkinson,Dryoxylononoharaense(Murayama),Euwallaceainterjectus(Blandford),Xyleborinusandrewesi(Blandford),Xyleborinusartestriatus(Eichhoff),Xyleborinusoctiesdentatus(Murayama),XyleborusbispinatusEichhoff,XyleborusseriatusBlandford,XyleborusspinulosusBlandford, andXylosandrusamputatus(Blandford).
Ambrosia beetles of the Euwallacea fornicatus (Eichhoff, 1868) species complex are emerging tree pests, responsible for significant damage to orchards and ecosystems around the world. The species complex comprises seven described species, all of which are nearly identical. Given that the morphology-defined species boundaries have been ambiguous, historically, there has been much disagreement on species validity, which was compounded by the presumed loss of the type series of E. fornicatus. The species complex was recently reviewed using morphometrics to associate the type specimens to the clades delineated with molecular data under the assumption of the lost type series. We rediscovered a syntype of Xyleborus fornicatus, and reevaluated the species in the complex using morphometrics. We propose the following taxonomic changes to the species complex: Euwallacea fornicatus (=E. tapatapaoensis (Schedl, 1951); = E. whitfordiodendrus (Schedl, 1942)) syn. res.); E. fornicatior (Eggers, 1923) (=E. schultzei (Schedl, 1951) syn. nov.); E. kuroshio (Gomez and Hulcr, 2018) and E. perbrevis (Schedl, 1951) stat. res. These taxonomic changes shift the species name associated with the widely used common names for two taxa, namely: Euwallacea fornicatus should be used for the “Polyphagous Shot Hole Borer”, and E. perbrevis for the “Tea Shot Hole Borer clade a”. A lectotype is designated for X. fornicatus in order to stabilize the use of the name.
1 November 2018 in Insect Systematics and Diversity
The ambrosia beetle Euwallacea fornicatus Eichhoff sensu lato is a complex of genetically divergent emerging pests responsible for damages to tree industries and ecosystems around the world. All lineages within the species complex are currently considered morphologically identical, presenting problems for their delineation and highlighting the shortcomings of species concepts based solely on type-specimen morphology. The objectives of this work were to 1) broaden the geographic sampling of the E. fornicatus complex in Asia, 2) reconstruct relationships between clades and populations, 3) find morphological characters or combinations of characters which are useful in delimiting the genetic lineages of the E. fornicatus species complex, and 4) propose taxonomic delimitation of species where morphology and phylogenetic identity correlate. Our integrated approach using molecular and morphological evidence suggests four clades that differ morphologically, but with overlap, therefore, cytochrome oxidase c subunit I (COI) barcoding remains necessary for assigning specimens to a clade. The following taxonomic changes are proposed: E. fornicatus (Eichhoff 1868) (= ‘Tea Shot Hole Borer Clade a’); E. fornicatior (Eggers 1923), stat. rev. (= ‘Tea Shot Hole Borer Clade b’); E. whitfordiodendrus (Schedl 1942), stat. rev. (= ‘Polyphagous Shot Hole Borer’); and E. kuroshio Gomez and Hulcr, sp. nov. (= ‘Kuroshio Shot Hole Borer’). This approach delivers a practical, evidence-based guidance for species delineation that can address overlapping variation in morphological characters of an emerging pest species complex.
Ambrosia beetles frequently invade non-native regions but are typically of no concern because most species live in dead trees and culture nonpathogenic symbiotic fungal gardens. Recently, however, several ambrosia beetle—fungus complexes have invaded non-native regions and killed large numbers of host trees. Such tree-killing invasions have occurred unexpectedly, and the mechanism of the ecological switch from dead trees to live trees has been left unexplained, or termed an “evolutionary mismatch.” We demonstrate that the mismatch hypothesis is not supported in the redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae), because this beetle is able to colonize live trees also in its native range and its symbiotic fungus acts as a pathogen in some native hosts. We further synthesized findings from recent literature and unpublished observations on several other invasive fungus-associated beetles such as Euwallacea fornicatus (Eggers) (Coleoptera: Curculionidae), Platypus qu... Resumen Los escarabajos descortezadores frecuentemente invaden las regiones no nativas, pero normalmente no son motivo de preocupación porque la mayoría de las especies viven en árboles muertos y cultivan hongos simbióticos no patógenos. Recientemente, sin embargo, varios complejos escarabajo descortezador-hongo han invadido regiones no nativas y han matado un gran número de árboles hospederos. Tales invasiones que matan árboles han ocurrido inesperadamente, y el mecanismo del cambio ecológico de árboles muertos a los árboles vivos se ha dejado inexplicado, o denominado un “desajuste evolutivo”. Demostramos que la hipótesis de desajuste no es apoyada en el escarabajo descortezador de aquacatillo, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae), porque este escarabajo es capaz de colonizar árboles vivos también en su área de distribución nativa y su hongo simbiótico actúa como patógeno en algunos hospederos nativos. Además, se sintetizaron los hallazgos de la literatura reciente y observaciones no...
Symbioses are increasingly seen as dynamic ecosystems with multiple associates and varying fidelity. Symbiont specificity remains elusive in one of the most ecologically successful and economically damaging eukaryotic symbioses: the ambrosia symbiosis of wood-boring beetles and fungi. We used multiplexed pyrosequencing of amplified internal transcribed spacer II (ITS2) ribosomal DNA (rDNA) libraries to document the communities of fungal associates and symbionts inside the mycangia (fungus transfer organ) of three ambrosia beetle species, Xyleborus affinis, Xyleborus ferrugineus and Xylosandrus crassiusculus. We processed 93 beetle samples from 5 locations across Florida, including reference communities. Fungal communities within mycangia included 14–20 fungus species, many more than reported by culture-based studies. We recovered previously known nutritional symbionts as members of the core community. We also detected several other fungal taxa that are equally frequent but whose function is unknown and many other transient species. The composition of fungal assemblages was significantly correlated with beetle species but not with locality. The type of mycangium appears to determine specificity: two Xyleborus with mandibular mycangia had multiple dominant associates with even abundances; Xylosandrus crassiusculus (mesonotal mycangium) communities were dominated by a single symbiont, Ambrosiella sp. Beetle mycangia also carried many fungi from the environment, including plant pathogens and endophytes. The ITS2 marker proved useful for ecological analyses, but the taxonomic resolution was limited to fungal genus or family, particularly in Ophiostomatales, which are under-represented in our amplicons as well as in public databases. This initial analysis of three beetle species suggests that each clade of ambrosia beetles and each mycangium type may support a functionally and taxonomically distinct symbiosis.