Multi-gene phylogeny of North American clear-winged moths (Lepidoptera: Sesiidae): a foundation for future evolutionary study of a speciose mimicry complex.

Sesiids are a diverse group of predominantly diurnal moths, many of which are Batesian mimics of Hymenoptera. However, their diversity and relationships are poorly understood. A multi-gene phylogenetic analysis of 48 North American sesiid species confirmed the traditional taxonomic tribal ranks, demonstrated the paraphyly of Carmenta and Synanthedon with respect to several other genera and ultimately provided minimal phylogenetic resolution within and between North American and European groups. Character support from each gene suggested inconsistency between the phylogenetic signal of the CAD gene and that of the other four genes. However, removal of CAD from subsequent phylogenetic analyses did not substantially change the initial phylogenetic results or return Carmenta and Synanthedon as reciprocally monophyletic, suggesting that it was not impacting the overall phylogenetic signal. The lack of resolution using genes that are typically informative at the species level for other lepidopterans suggests a surprisingly rapid radiation of species in Carmenta/Synanthedon. This group also exhibits a wide range of mimicry strategies and hostplant usage, which could be fertile ground for future study.

Patterns of host tree use within a lineage of saproxlic snout-less weevils (Coleoptera: Curculionidae: Scolytinae: Scolytini).

The influence of plants in the diversification of herbivorous insects, specifically those that utilize moribund and dead hosts, is little explored. Host shifts are expected because the effectiveness of toxic secondary chemicals is lessened by decay of dead plants. Feeding on dead plants also releases herbivorous insect lineages from diversifying within a particular plant lineage. Thus, phylogenetic constraints on the herbivorous insect lineage imposed by the host plants are diminished and repeated patterns of species diversification in an association with unrelated host trees is hypothesized (i.e., taxon cycle). Scolytini, a diverse weevil tribe, specialize on many different dead and moribund plant taxa as a source of food. These species and their hosts offer an opportunity to examine the association between dead host plants and the extent of phylogenetic constraints. A phylogeny of the Scolytini was reconstructed with likelihood and Bayesian analyses of DNA sequence data from nuclear (28S, CAD, ArgK) and mitochondrial (COI) genes. Ancestral host usage and geography was reconstructed using likelihood criteria and conservation of host use was tested. Results supported a monophyletic Scolytini, Ceratolepis, Loganius, and a paraphyletic Scolytus, Camptocerus and Cnemonyx. Diversification of the Scolytini generally occurred well after their host taxa diversified and suggests a sequential evolution of host use. In this scenario the beetle imposes little selection pressure on the tree but the tree provides a platform for beetle evolution. Major changes in host tree use occurred during periods of global cooling associated with changes in beetle biogeography. Diversification of beetles occurred on common and widespread hosts and there was likely a single origination of conifer-feeding from angiosperm-feeding species during the early Pliocene and a radiation of beetle species from the Palearctic to the Nearctic. Overall, the observed patterns of Scolytini host use are conserved and are similar to those expected in a taxon pulse diversification. That is, after a host switch to an unrelated tree, the beetles diversify within the host plant lineage. The need to locate an ephemeral food resource, i.e., a dying tree, likely maintains host specificity once a host shift occurs. These findings suggest that characteristics of dead and moribund host plants (e.g. secondary chemicals) influence the diversification of these saproxlic weevils despite the reduction of selection pressures.

A highly resolved food web for insect seed predators in a species-rich tropical forest.

The top-down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species-rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co-occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host-specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen-Connell mechanism.

Phylogenomics clarifies repeated evolutionary origins of inbreeding and fungus farming in bark beetles (Curculionidae, Scolytinae).

Bark and ambrosia beetles (Curculionidae, Scolytinae) display a conspicuous diversity of unusual genetic and ecological attributes and behaviors. Reconstructing the evolution of Scolytinae, particularly the large and ecologically significant tribe Cryphalini (pygmy borers), has long been problematic. These challenges have not adequately been addressed using morphological characters, and previous research has used only DNA sequence data from small numbers of genes. Through a combination of anchored hybrid enrichment, low-coverage draft genomes, and transcriptomes, we addressed these challenges by amassing a large molecular phylogenetic dataset for bark and ambrosia beetles. The resulting DNA sequence data from 251 protein coding genes (114,276 bp of nucleotide sequence data) support inference of the first robust phylogeny of Scolytinae, with a special focus on the species rich tribe Cryphalini and its close relatives. Key strategies, including inbreeding mating systems and fungus farming, evolved repeatedly across Scolytinae. We confirm 12 of 16 hypothesized origins of fungus farming, 6 of 8 origins of inbreeding polygyny and at least 11 independent origins of a super-generalist host range. These three innovations are statistically correlated, but their appearance within lineages was not necessarily simultaneous. Additionally, the evolution of extreme host plant generalism often preceded, rather than succeeded, fungus farming. Of the high-diversity tribes of Scolytinae, only Xyleborini is monophyletic, Corthylini is paraphyletic and Cryphalini is highly polyphyletic. Cryphalini sensu stricto is part of a clade containing the genera Hypothenemus, Cryphalus and Trypophloeus, and the tribe Xyloterini. Stegomerus and Cryptocarenus (Cryphalini) are part of a clade otherwise containing all Corthylini. Several other genera, including Ernoporus and Scolytogenes (Cryphalini), make up a distantly related clade. Several of the genera of Cryphalini are also intermixed. For example, Cryphalus and Hypocryphalus are intermingled, as well as Ernoporicus, Ptilopodius and Scolytogenes. Our data are consistent with widespread polyphyly and paraphyly across Scolytinae and within Cryphalini, and provides new insights into the evolution of inbreeding mating systems and fungus farming in the species rich and ecologically significant weevil subfamily Scolytinae.

PCR Multiplexes Discriminate Fusarium Symbionts of Invasive Euwallacea Ambrosia Beetles that Inflict Damage on Numerous Tree Species Throughout the United States.

Asian Euwallacea ambrosia beetles vector Fusarium mutualists. The ambrosial fusaria are all members of the ambrosia Fusarium clade (AFC) within the Fusarium solani species complex (FSSC). Several Euwallacea-Fusarium mutualists have been introduced into nonnative regions and have caused varying degrees of damage to orchard, landscape, and forest trees. Knowledge of symbiont fidelity is limited by current identification methods, which typically requires analysis of DNA sequence data from beetles and the symbionts cultured from their oral mycangia. Here, polymerase chain reaction (PCR)-based diagnostic tools were developed to identify the six Fusarium symbionts of exotic Euwallacea spp. currently known within the United States. Whole-genome sequences were generated for representatives of six AFC species plus F. ambrosium and aligned to the annotated genome of F. euwallaceae. Taxon-specific primer-annealing sites were identified that rapidly distinguish the AFC species currently within the United States. PCR specificity, reliability, and sensitivity were validated using a panel of 72 Fusarium isolates, including 47 reference cultures. Culture-independent multiplex assays accurately identified two AFC fusaria using DNA isolated from heads of their respective beetle partners. The PCR assays were used to show that Euwallacea validus is exclusively associated with AF-4 throughout its sampled range within eastern North America. The rapid assay supports federal and state agency efforts to monitor spread of these invasive pests and mitigate further introductions.

The evolution of Batesian mimicry within the North American Asidini (Coleoptera: Tenebrionidae).

The asidine darkling beetles (Coleoptera: Tenebrionidae: Asidini) are a diverse tribe of flightless tenebrionids found in many arid and sub-arid habitats around the world. The 263 currently described North American species are contained in ten genera, all of which are restricted to the western half of the continent. The Asidini, like all members of the subfamily Pimeliinae, lack defensive glands. Instead, several phenotypic traits occur within the tribe that may help limit predation. These include the contrasting defensive strategies of crypsis, through either background matching or pattern disruption, and Batesian mimicry of the chemically defended genus Eleodes. Dorsal elytral morphology was assessed between 53 North American asidine species and 13 common Eleodes model species using multiple methodologies to assess similarities between species in the two groups that might indicate mimetic relationships. A phylogeny of the North American asidines is used to map the occurrence of differing defensive strategies within the tribe. Crypsis is reconstructed as the ancestral state, with two origins for Batesian mimicry and multiple reversals. The combination of strongly to weakly cryptic species and varying levels of mimetic fidelity to Eleodes model species make the asidines a promising lineage upon which to further explore the evolution of defensive phenotypes.

History of the Exotic Ambrosia Beetles Euwallacea interjectus and Euwallacea validus (Coleoptera: Curculionidae: Xyleborini) in the United States.

Exotic insects are constantly intercepted at U.S. ports-of-entry. Of these, wood-boring beetles, particularly xyleborine ambrosia beetles, are sometimes missed during port inspections and become established in the United States. Euwallacea validus (Eichhoff) and Euwallacea interjectus (Blandford) are morphologically similar Asian ambrosia beetle species that vary by their fungal associates and their potential to cause economic damage. Euwallacea validus and E. interjectus were first discovered in New York (1975) and Hawaii (1976), respectively. Euwallacea validus was collected multiple times from widely separated localities and is assumed to have spread throughout the eastern United States. The discovery of E. interjectus in Florida (2011) and Texas (2011) prompted our review of the E. validus specimens because of the potential misidentification of the species. In addition, using mitochondrial cytochrome oxidase I (COI) DNA data and phylogenetic analysis, we tested the hypothesis that multiple introductions account for the U.S. populations of E. interjectus and E. validus. Our review of 7,184 specimens revealed an earlier introduction to the mainland for E. interjectus, which was first collected from Louisiana in 1984. This species is distributed in the South while E. validus occurs in the North with a known area of syntopy in northeastern Georgia. The extent of the syntopy within the United States is unknown and further investigation is required. Phylogenetic analysis of 24 E. interjectus and 20 E. validus individuals resolved clades that associated with each species and gross geographic provenance. Four well-supported clades represented E. interjectus which included the following localities: 1) Hawaii and Thailand; 2) Vietnam, Taiwan, and Texas; 3) Okinawa (Japan); and 4) Japan and several southern U.S. states. One clade comprised all E. validus specimens from Japan and the mainland United States. Four and two haplotypes were found for the E. interjectus and E. validus specimens, respectively, in mainland United States. Except for the Texas specimen, the haplotypes differed by one nucleotide. The relationship of the haplotypes and their sequence similarity suggested that the provenance of E. validus and the majority of E. interjectus haplotypes was Japan while the Texas haplotype originated later and from a location near Taiwan. Given the high nucleotide sequence difference between the Hawaiian and Thai haplotypes, the exact origin of the Hawaiian E. interjectus is unknown but likely Southeast Asia. A broader investigation including more SE Asian individuals will help to further explain the introduction of E. interjectus into Hawaii and Texas.

Coptoborus ochromactonus, n. sp. (Coleoptera: Curculionidae: Scolytinae), an emerging pest of cultivated balsa (Malvales: Malvaceae) in Ecuador.

A new species of xyleborine ambrosia beetle has been found to attack balsa, Ochroma pyramidale (Cavanilles ex Lamarck) Urban, in Ecuador. Coptoborus ochromactonus Smith & Cognato is described and its biology is reported. Large-scale surveys were conducted between 2006 and 2009, and observational studies were carried out between 2010 and 2013 in Ecuadorian commercial plantations to determine life history and host preference characteristics. C. ochromactonus attacked balsa between 1.5 and 3 yr in age. Successful attacks were more prevalent in smaller diameter trees and unhealthy trees. In general, attacks and beetle-caused mortality were more prevalent during the dry summer months when trees were under more moisture and light stress. Fungal mycelia were consistently observed coating beetle galleries and are likely the true damaging agent to balsa trees.

 Tricosauniseriata, a new species of xyleborine ambrosia beetle from Thailand (Coleoptera, Curculionidae, Scolytinae, Xyleborini).

A new species, Tricosauniseriatasp. nov., is described here. A list of Tricosa species found in Thailand with distributions and an updated key to Tricosa are also provided.

Fusarium kuroshium is the primary fungal symbiont of an ambrosia beetle, Euwallacea fornicatus, and can kill mango tree in Japan.

This study identifies fungi associated with Euwallacea fornicatus and determines whether these fungal species play the role of primary symbiont. E. fornicatus adults that emerged from the branches of infested trees in Okinawa main island, Japan, were collected and used to isolate fungi. Fusarium kuroshium and Penicillium citrinum were the most dominant fungal associates of females and males, respectively. F. kuroshium was much more frequently isolated from the head, including mycangia (fungus-carrying organs), of females than any other body parts. We inoculated healthy mango saplings with F. kuroshium or F. decemcellulare, both of which were symbionts of E. fornicatus females infesting mango trees. F. kuroshium decreased leaf stomatal conductance and rate of xylem sap-conduction area and increased length and area of xylem discoloration of the saplings, thereby weakening and killing some. These results suggest that F. kuroshium, a mycangial fungus of E. fornicatus, inhibits water flow in mango trees. This study is the first to report that F. kuroshium causes wilt disease in mango trees and that it is a primary fungal symbiont of E. fornicatus.

Molecular phylogeny of bark and ambrosia beetles reveals multiple origins of fungus farming during periods of global warming.

BACKGROUND: Fungus farming is an unusual life style in insects that has evolved many times in the wood boring weevils named 'ambrosia beetles'. Multiple occurrences of this behaviour allow for a detailed comparison of the different origins of fungus farming through time, its directionality, and possible ancestral states. We tested these hypotheses with a phylogeny representing the largest data set to date, nearly 4 kb of nucleotides from COI, EF-1α, CAD, ArgK, 28S, and 200 scolytine taxa. RESULTS: Phylogenetic analyses using Bayesian or parsimony approaches placed the root of Scolytinae close to the tribe Scolytini and Microborus, but otherwise indicated low resolution at older nodes. More recent clades were well resolved, including ten origins of fungus farming. There were no subsequent reversals to bark or phloem feeding in the fungus farming clades. The oldest origin of fungus farming was estimated near 50 Ma, long after the origin of Scolytinae (100-120 Ma). Younger origins included the species rich Xyleborini, dated to 21 Ma. Sister group comparisons and test of independence between traits indicated that neither gregarious larval feeding nor regular inbreeding by sibling mating was strongly correlated with the origin of fungus farming. CONCLUSION: Origins of fungus farming corresponded mainly with two periods of global warming in the Cenozoic era, which were characterised by broadly distributed tropical forests. Hence, it seems likely that warm climates and expanding tropical angiosperm forests played critical roles in the successful radiation of diverse fungus farming groups. However, further investigation will likely reveal additional biological factors that promote fungus farming.

Ecuadorian Coptoborus beetles harbor Fusarium and Graphium fungi previously associated with Euwallacea ambrosia beetles.

Ambrosia beetles from the scolytine tribe Xyleborini (Curculionidae) are important to the decomposition of woody plant material on every continent except Antarctica. These insects farm fungi on the walls of tunnels they build inside recently dead trees and rely on the fungi for nutrition during all stages of their lives. Such ambrosia fungi rely on the beetles to provide appropriate substrates and environmental conditions for growth. A small minority of xyleborine ambrosia beetle-fungal partnerships cause significant damage to healthy trees. The xyleborine beetle Coptoborus ochromactonus vectors a Fusarium (Hypocreales) fungus that is lethal to balsa (Ochroma pyramidale (Malvaceae)) trees in Ecuador. Although this pathogenic fungus and its associated beetle are not known to be established in the United States, several other non-native ambrosia beetle species are vectors of destructive plant diseases in this country. This fact and the acceleration of trade between South America and the United States demonstrate the importance of understanding fungal plant pathogens before they escape their native ranges. Here we identify the fungi accompanying Coptoborus ambrosia beetles collected in Ecuador. Classification based ribosomal internal transcribed spacer 1 (ITS) sequences revealed the most prevalent fungi associated with Coptoborus are Fusarium sp. and Graphium sp. (Microascales: Microascaceae), which have been confirmed as ambrosia fungi for xyleborine ambrosia beetles, and Clonostsachys sp. (Hypocreales), which is a diverse genus found abundantly in soils and associated with plants. Phylogenetic analyses of the Fusarium strains based on ITS, translation elongation factor (EF1-α), and two subunits of the DNA-directed RNA polymerase II (RPB1 and RPB2) identified them as Fusarium sp. AF-9 in the Ambrosia Fusarium Clade (AFC). This Fusarium species was previously associated with a few xyleborine ambrosia beetles, most notably the species complex Euwallacea fornicatus (Eichhoff 1868) (Curculionidae: Scolytinae: Xyleborini). Examination of ITS and EF1-α sequences showed a close affinity between the Graphium isolated from Coptoborus spp. and other xyleborine-associated Graphium as well as the soil fungus Graphium basitruncatum. This characterization of ambrosia fungi through DNA sequencing confirms the identity of a putative plant pathogen spread by Coptoborus beetles and expands the documented range of Fusarium and Graphium ambrosia fungi.

One hundred eighteen taxonomic changes among Xyleborine ambrosia beetles (Coleoptera: Curculionidae: Scolytinae).

An ongoing study of the ambrosia beetle tribe Xyleborini has resulted in numerous taxonomic changes mostly representing new generic/species combinations which remove species from the once all-encompassing Xyleborus Eichhoff, 1864 to other genera based on revised taxonomic concepts. These changes are here listed. Terminalinus Hopkins, 1915 is removed from synonymy with Cyclorhipidion Hagedorn, 1912 and reinstated as a valid genus. Five species are removed from synonymy and reinstated as valid species: Amasa brevipennis (Schedl, 1971), Amasa fulgens (Schedl, 1975), Ambrosiophilus immitatrix (Schedl, 1975), Ambrosiophilus semirufus (Schedl, 1959), Microperus leprosulus (Schedl, 1936). The following 97 new or restored combinations are proposed: Ambrosiophilus bispinosulus (Schedl, 1961) comb. nov., Ambrosiophilus compressus (Lea, 1894) comb. nov., Ambrosiophilus latecompressus (Schedl, 1936) comb. nov., Ambrosiophilus pertortuosus (Schedl, 1942) comb. nov., Ambrosiophilus tomicoides (Eggers, 1923) comb. nov., Ambrosiophilus tortuosus (Schedl, 1942) comb. nov., Euwallacea obliquecauda (Motschulsky, 1863) comb. nov., all from Ambrosiodmus Hopkins, 1915; Coptodryas decepta (Schedl, 1979) comb. nov., Microperus pusillus (Eggers, 1927) comb. nov., both from Arixyleborus Hopkins, 1915; Coptodryas pseudopunctula (Schedl, 1942) comb. nov., from Cnestus Sampson, 1911; Microperus abbreviatus (Schedl, 1942) comb. nov., Microperus amphicauda (Browne, 1986) comb. nov., Microperus borneensis (Browne, 1986) comb. nov., Microperus comptus (Sampson, 1919) comb. nov., Microperus gorontalosus (Schedl, 1939) comb. nov., Microperus pullus (Schedl, 1952) comb. nov., Microperus tenellus (Schedl, 1959) comb. nov., Microperus vafer Schedl, 1957 comb. nov., all from Coptodryas Hopkins, 1915; Ambrosiophilus pityogenes (Schedl, 1936) comb. nov., Arixyleborus scapularis (Schedl, 1942) comb. nov., Beaverium dihingicum (Wood, 1992) comb. nov., Beaverium rufonitidus (Schedl, 1951) comb. nov., Coptodryas brevior (Eggers) comb. nov., Terminalinus dipterocarpi Hopkins, 1915 comb. res., Terminalinus sexspinatus (Schedl, 1935) comb. nov., Terminalinus terminaliae (Hopkins, 1915) comb. res., Truncaudum leverensis (Browne, 1986) comb. nov., all from Cyclorhipidion Hagedorn, 1912; Planiculus kororensis (Wood, 1960) comb. nov., Planiculus loricatus (Schedl, 1933) comb. nov., Planiculus murudensis (Browne, 1965) comb. nov., all from Euwallacea Reitter, 1915; Terminalinus anisopterae (Browne, 1983) comb. nov., Terminalinus indigens (Schedl, 1955) comb. nov., Terminalinus macropterus (Schedl, 1935) comb. nov., Terminalinus major (Stebbing, 1909) comb. nov., Terminalinus pilifer (Eggers, 1923) comb. nov., Terminalinus posticepilosus (Schedl, 1951) comb. res., Terminalinus pseudopilifer (Schedl, 1936) comb. nov., Terminalinus sulcinoides (Schedl, 1974) comb. nov., all from Fortiborus Hulcr & Cognato, 2010; Microperus micrographus (Schedl, 1958) comb. nov., Microperus truncatipennis (Schedl, 1961) comb. nov., both from Xyleborinus Reitter, 1913; Ambrosiophilus immitatrix (Schedl, 1975) comb. nov., Ambrosiophilus semirufus (Schedl, 1959) comb. nov., Arixyleborus crenulatus (Eggers, 1920) comb. nov., Arixyleborus strombosiopsis (Schedl, 1957) comb. nov., Beaverium batoensis (Eggers, 1923) comb. nov., Beaverium calvus (Schedl, 1942) comb. nov., Beaverium obstipus (Schedl, 1935) comb. nov., Beaverium rufus (Schedl, 1951) comb. nov., Coptodryas cuneola (Eggers, 1927) comb. nov., Cyclorhipidion amanicum (Hagedorn, 1910) comb. nov., Cyclorhipidion impar (Eggers, 1927) comb. nov., Cyclorhipidion inaequale (Schedl, 1934) comb. nov., Cyclorhipidion kajangensis (Schedl, 1942) comb. nov., Cyclorhipidion obiensis (Browne, 1980) comb. nov., Cyclorhipidion obtusatum (Schedl, 1972) comb. nov., Cyclorhipidion perpunctatum (Schedl, 1971) comb. nov., Cyclorhipidion repositum (Schedl) comb. nov., Cyclorhipidion separandum (Schedl, 1971) comb. nov., Debus abscissus (Browne, 1974) comb. nov., Debus amplexicauda (Hagedorn, 1910) comb. nov., Debus armillatus (Schedl, 1933) comb. nov., Debus balbalanus (Eggers 1927) comb. nov., Debus blandus (Schedl, 1954) comb. nov., Debus cavatus (Browne, 1980) comb. nov., Debus cylindromorphus (Eggers, 1927) comb. nov., Debus dentatus (Blandford, 1895) comb. nov., Debus excavus (Schedl, 1964) comb. nov., Debus fischeri (Hagedorn, 1908) comb. nov., Debus hatanakai (Browne, 1983) comb. nov., Debus insitivus (Schedl, 1959) comb. nov., Debus persimilis (Eggers, 1927) comb. nov., Debus subdentatus (Browne, 1974) comb. nov., Debus trispinatus (Browne, 1981) comb. nov., Diuncus taxicornis (Schedl, 1971) comb. nov., Euwallacea agathis (Browne, 1984) comb. nov., Euwallacea assimilis (Eggers, 1927) comb. nov., Euwallacea bryanti (Sampson, 1919) comb. nov., Euwallacea latecarinatus (Schedl, 1936) comb. nov., Euwallacea pseudorudis (Schedl, 1951) comb. nov., Euwallacea semipolitus (Schedl, 1951) comb. nov., Euwallacea temetiuicus (Beeson, 1935) comb. nov., Immanus duploarmatus (Browne, 1962) comb. nov., Leptoxyleborus sublinearis (Eggers, 1940) comb. nov., Peridryocoetes pinguis (Browne, 1983) (Dryocoetini) comb. nov., Stictodex halli (Schedl, 1954) comb. nov., Stictodex rimulosus (Schedl, 1959) comb. nov., Terminalinus granurum (Browne, 1980) comb. nov., Terminalinus indonesianus (Browne, 1984) comb. nov., Terminalinus moluccanus (Browne, 1985) comb. nov., Terminalinus pseudomajor (Schedl, 1951) comb. nov., Terminalinus sublongus (Eggers, 1927) comb. nov., Terminalinus takeharai (Browne) comb. nov., Terminalinus xanthophyllus (Schedl, 1942) comb. res., Tricosa abberrans (Schedl, 1959) comb. nov., Xenoxylebora truncatula (Schedl, 1957) comb. nov., Xyleborinus figuratus (Schedl, 1959) comb. nov., Xylosandrus cancellatus (Eggers, 1936) comb. nov., all from Xyleborus. Fifteen new synonyms are proposed: Anisandrus ursulus (Eggers, 1923)(= Xyleborus lativentris Schedl, 1942 syn. nov.); Cyclorhipidion amanicus (Hagedorn, 1910)(= Xyleborus jongaensis Schedl, 1941 syn. nov.); Cyclorhipidion bodoanum (Reitter, 1913) (= Xyleborus takinoyensis Murayama, 1953 syn. nov.); Cyclorhipidion pelliculosum (Eichhoff, 1878) (= Xyleborus okinosenensis Murayama, 1961 syn. nov.); Cyclorhipidion repositum (Schedl, 1942) (= Xyleborus pruinosulus Browne, 1979 syn. nov.); Debus persimilis (Eggers, 1927) (= Xyleborus subdolosus Schedl, 1942c syn. nov.); Debus robustipennis (Schedl, 1954) (= Xyleborus interponens Schedl, 1954 syn. nov.); Euwallacea destruens (Blandford, 1896) (= Xyleborus procerior Schedl, 1942 syn. nov.); Euwallacea nigrosetosus (Schedl, 1939) (= Xyleborus nigripennis Schedl, 1951 syn. nov.); Euwallacea siporanus (Hagedorn, 1910) (= Xyleborus perakensis Schedl, 1942 syn. nov.); Microperus quercicola (Eggers, 1926) (= Xyleborus semistriatus Schedl, 1971 syn. nov.); Stictodex dimidiatus (Eggers, 1927) (= Xyleborus spicatus Browne, 1986 syn. nov.); Stictodex halli (Schedl, 1954) (= Xyleborus cuspidus Schedl, 1975 syn. nov.); Terminalinus Hopkins, 1915 (= Fortiborus Hulcr & Cognato 2010 syn. nov.); Terminalinus moluccanus (Browne, 1985) (= Xyleborus teminabani Browne, 1986 syn. nov.).

New species and new records of Xyleborini from the Oriental region, Japan and Papua New Guinea (Coleoptera: Curculionidae: Scolytinae).

Eighteen xyleborine ambrosia beetles are described and illustrated: Anisandrus proscissus Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Anisandrus simplex Smith, Beaver & Cognato sp. nov. (Nepal), Arixyleborus belalongi Smith, Beaver & Cognato sp. nov. (Brunei Darussalam), Beaverium brevicaudatus Smith, Beaver & Cognato sp. nov. (Indonesia), Cnestus luculentus Smith, Beaver & Cognato sp. nov. (India), Cyclorhipidion achlys Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Cyclorhipidion conidentatus Smith, Beaver & Cognato sp. nov. (Indonesia), Cyclorhipidion gladigerum Smith, Beaver & Cognato sp. nov. (Thailand), Cyclorhipidion lapilliferum Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Cyclorhipidion nepalense Smith, Beaver & Cognato sp. nov. (Nepal), Cyclorhipidion taedulum Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Cyclorhipidion titorum Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Euwallacea alastos Smith, Beaver & Cognato sp. nov. (Japan), Leptoxyleborus regina Smith, Beaver & Cognato sp. nov. (Papua New Guinea), Tricosa hipparion Smith, Beaver & Cognato sp. nov. (Malaysia), Xyleborinus acanthopteron Smith, Beaver & Cognato sp. nov. (Thailand), Xyleborinus dumosus Smith, Beaver, Pham & Cognato sp. nov. (Vietnam), Xyleborinus nobuchii Smith, Beaver & Cognato sp. nov. (Japan). New distribution records are reported for 67 Asian species. Cyclorhipidion nemesis Smith & Cognato, described from U. S. A., is reported from Asia (China), its hypothesized native continent, for the first time. Its identity is confirmed with COI and CAD DNA within a phylogenetic analysis including other Cyclorhipidion species.

The age and phylogeny of wood boring weevils and the origin of subsociality.

A large proportion of the hyperdiverse weevils are wood boring and many of these taxa have subsocial family structures. The origin and relationship between certain wood boring weevil taxa has been problematic to solve and hypotheses on their phylogenies change substantially between different studies. We aimed at testing the phylogenetic position and monophyly of the most prominent wood boring taxa Scolytinae, Platypodinae and Cossoninae, including a range of weevil outgroups with either the herbivorous or wood boring habit. Many putatively intergrading taxa were included in a broad phylogenetic analysis for the first time in this study, such as Schedlarius, Mecopelmus, Coptonotus, Dactylipalpus, Coptocorynus and allied Araucariini taxa, Dobionus, Psepholax, Amorphocerus-Porthetes, and some peculiar wood boring Conoderini with bark beetle behaviour. Data analyses were based on 128 morphological characters, rDNA nucleotides from the D2-D3 segment of 28S, and nucleotides and amino acids from the protein encoding gene fragments of CAD, ArgK, EF-1α and COI. Although the results varied for some of the groups between various data sets and analyses, one may conclude the following from this study: Scolytinae and Platypodinae are likely sister lineages most closely related to Coptonotus; Cossoninae is monophyletic (including Araucariini) and more distantly related to Scolytinae; Amorphocerini is not part of Cossoninae and Psepholax may belong to Cryptorhynchini. Likelihood estimation of ancestral state reconstruction of subsociality indicated five or six origins as a conservative estimate. Overall the phylogenetic results were quite dependent on morphological data and we conclude that more genetic loci must be sampled to improve phylogenetic resolution. However, some results such as the derived position of Scolytinae were consistent between morphological and molecular data. A revised time estimation of the origin of Curculionidae and various subfamily groups were made using the recently updated fossil age of Scolytinae (100 Ma), which had a significant influence on node age estimates.

A revision of the Neotropical genus Coptoborus Hopkins (Coleoptera, Curculionidae, Scolytinae, Xyleborini).

The Neotropical xyleborine ambrosia beetle genus Coptoborus Hopkins is reviewed. The following 40 Coptoborus species are described: C. amplissimus sp. nov. (Peru), C. asperatus sp. nov. (Ecuador), C. barbicauda sp. nov. (French Guiana), C. bettysmithae sp. nov. (Ecuador), C. brevicauda sp. nov. (Ecuador), C. brigman sp. nov. (Ecuador), C. busoror sp. nov. (Ecuador), C. capillisoror sp. nov. (Brazil), C. chica sp. nov. (Suriname), C. crassisororcula sp. nov. (Peru), C. doliolum sp. nov. (Ecuador), C. erwini sp. nov. (Ecuador), C. furiosa sp. nov. (Ecuador), C. galacatosae sp. nov. (Ecuador), C. hansen sp. nov. (Brazil), C. incomptus sp. nov. (Peru), C. janeway sp. nov. (Peru), C. katniss sp. nov. (Ecuador), C. leeloo sp. nov. (Ecuador), C. leia sp. nov. (Ecuador, Suriname), C. leporinus sp. nov. (Peru), C. martinezae sp. nov. (Ecuador), C. murinus sp. nov. (Ecuador), C. newt sp. nov. (Peru), C. osbornae sp. nov. (Ecuador), C. panosus sp. nov. (French Guiana), C. papillicauda sp. nov. (Suriname), C. pilisoror sp. nov. (Ecuador), C. ripley sp. nov. (Ecuador), C. sagitticauda sp. nov. (Guyana), C. sarahconnor sp. nov. (Brazil), C. scully sp. nov. (Ecuador), C. sicula sp. nov. (Ecuador), C. sororcula sp. nov. (Peru), C. starbuck sp. nov. (Ecuador), C. trinity sp. nov. (Brazil), C. uhura sp. nov. (Peru), C. vasquez sp. nov. (Panama), C. vrataski sp. nov. (Brazil), and C. yar sp. nov. (Ecuador). Seventeen new combinations are given: Coptoborus amazonicus (Petrov, 2020) comb. nov., C. atlanticus (Bright & Torres, 2006) comb. nov., C. bellus Bright & Torres, 2006 comb. nov., C. coartatus (Sampson, 1921) comb. nov., C. crinitulus (Wood, 1974) comb. nov., C. exilis (Schedl, 1934) comb. nov., C. incultus (Wood, 1975) comb. nov., C. magnus (Petrov, 2020) comb. nov., C. micarius (Wood, 1974) comb. nov., C. obtusicornis (Schedl, 1976) comb. nov., C. paurus (Wood, 2007) comb. nov., C. pristis (Wood, 1974) comb. nov., C. pseudotenuis (Schedl, 1936) comb. nov., C. puertoricensis (Bright & Torres, 2006) comb. nov., C. ricini (Eggers, 1932) comb. nov., C. semicostatus (Schedl, 1948) comb. nov., C. tristiculus (Wood, 1975) comb. nov., and C. villosulus (Blandford, 1898) comb. nov. Two new synonyms are proposed: Coptoborus Hopkins, 1915 (= Theoborus Hopkins, 1915 syn. nov.) and Coptoborus villosulus (Blandford, 1898) (= Theoborus theobromae Hopkins, 1915 syn. nov.). Xyleborus neosphenos Schedl, 1976 comb. res. is removed from Coptoborus. The revised genus now contains 77 species and a key to their identification is provided.

Species-rich bark and ambrosia beetle fauna (Coleoptera, Curculionidae, Scolytinae) of the Ecuadorian Amazonian Forest Canopy.

Canopy fogging was used to sample the diversity of bark and ambrosia beetles (Coleoptera, Curculionidae, Scolytinae) at two western Amazonian rainforest sites in Ecuador. Sampling was conducted by Dr Terry Erwin and assistants from 1994-2006 and yielded 1158 samples containing 2500 scolytine specimens representing more than 400 morphospecies. Here, we analyze a subset of these data representing two ecological groups: true bark beetles (52 morphospecies) and ambrosia beetles (69 morphospecies). A high percentage of these taxa occurred as singletons and doubletons and their species accumulation curves did not reach an asymptote. Diversity estimates placed the total scolytine species richness for this taxon subset present at the two sites between 260 and 323 species. The α-diversity was remarkably high at each site, while the apparently high ß-diversity was an artifact of undersampling, as shown by a Monte Carlo resampling analysis. This study demonstrates the utility of canopy fogging for the discovery of new scolytine taxa and for approximate diversity assessment, but a substantially greater sampling effort would be needed for conclusive alpha as well as beta diversity estimates.

Polyphyly of Xylosandrus Reitter inferred from nuclear and mitochondrial genes (Coleoptera: Curculionidae: Scolytinae).

The Xyleborina ambrosia beetle genus Xylosandrus contains 54 species, several of which are of economic importance. The monophyly of the genus was tested using a data set comprised of multiple gene loci: 28S rDNA; the mitochondrial gene cytochrome oxidase I (COI); and the nuclear genes arginine kinase (ArgK), rudimentary (CAD), and Elongation Factor 1alpha (EF-1alpha). The nuclear protein-coding genes CAD and ArgK were used for the first time in phylogenetics of Scolytinae. Analyses were performed using Parsimony and Bayesian optimality criteria. Our analyses included 43 specimens representing 15 Xylosandrus species and 20 species from Amasa, Anisandrus, Cnestus, Euwallacea and Xyleborus, and two species from the outgroup genus Coccotrypes. All analyses recovered a polyphyletic Xylosandrus. Several species of Xylosandrus were consistently placed in clades with the genera Anisandrus and Cnestus with high support values (100% bootstrap support). Among these, was the economically important invasive species X. mutilatus, which was consistently recovered as part of the "Cnestus" clade. In our analyses, both CAD and ArgK demonstrated phylogenetic utility across varying nodal depths. Despite the selection of genes with signals at complementary phylogenetic depths, the data set used herein did not resolve the phylogeny of Xylosandrus and related genera. Since the taxon sample available for molecular work represents only a fraction of Xylosandrus species, a complete revision that combines molecular and morphological data in a total evidence approach is recommended for the genus.

DNA identification confirms pecan weevil (Coleoptera: Curculionidae) infestation of Carpathian walnut.

Larvae found infesting fruit from a Carpathian walnut, Juglans regia L., tree in Missouri were confirmed by DNA analysis to be those of pecan weevil, Curculio caryae (Horn) (Coleoptera: Curculionidae). The infested walnut tree occurs in the midst of pecan weevil-infested pecans, Carya illinoinensis (Wang.) K. Koch; the larval haplotypes were found to be identical to pecan weevil larvae from the region, indicating that the walnut infestation arose by association with infested pecan. This is the first confirmed DNA analysis showing pecan weevil attacks J. regia and the second report that J. regia may be at risk of infestation by pecan weevil. Further study indicates this infestation on walnut is established and ongoing. The pecan weevil is a key pest of pecan and seems capable of inflicting similar damage to walnut if spread to commercial areas that produce J. regia.

Species composition, seasonal activity, and semiochemical response of native and exotic bark and ambrosia beetles (Coleoptera: Curculionidae: Scolytinae) in northeastern Ohio.

In 2007, we surveyed the alien and endemic scolytine (bark and ambrosia beetles) fauna of northeastern Ohio, and for the most abundant species, we characterized their seasonal activity and response to three semiochemical baits. In total ,5,339 scolytine beetles represented by 47 species and 29 genera were caught in Lindgren funnel traps. Three species constituted 57% of the total catch, including Xylosandrus germanus (Blandford), Tomicus piniperda (L.), and Dryocoetes autographus (Ratzeburg). Of the total captured, 32% of the species and approximately 60% of the individuals were exotic, suggesting that exotic species numerically dominate the scolytine fauna in some urban areas. More native and exotic species were caught in traps baited with ethanol alone than in traps baited with other lures. However, significantly more individuals, especially of T. piniperda, D. autographus, Gnathotrichus materiarius (Fitch), and Ips grandicollis (Eichhoff), and species were caught in traps baited with ethanol plus alpha-pinene than in traps baited with ethanol alone or the exotic Ips lure. This suggests that among these baits, the ethanol plus alpha-pinene baits may be useful in maximizing scolytine beetle catches of these species within this region. Species diversity and richness for both native and exotic beetles was greatest in traps baited with ethanol alone. The period of peak trap capture varied depending upon species: X. germanus was most abundant in traps in mid-May and early-August; T. piniperda in mid-May; D. autographus in early June, mid-July, and mid-September; Anisandrus sayi Hopkins and G. materiarius in mid-May, mid-July, and early September; and I. grandicollis in early April, mid-July, and late September.