Anthony I. Cognato

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

Abstract Ambrosia-feeding scolytids in the Xyleborina are the most abundant exotic bark beetles in North America. Many new species discoveries and taxonomic changes have occurred since their last review in 1982. The need for a worldwide revision of this tribe hampers the ability to identify species. To remedy this situation, an illustrated key is constructed and Xyleborus glabratus Eichhoff, Xyleborus similis Ferrari, and Euwallacea fornicatus (Eichhoff), which are new to North America, are reported and diagnosed. In addition the key is presented in electronic format with additional illustration (http://scolytid.msu.edu) to increase worldwide availability and accommodate future taxonomic revision.

Standard Percent DNA Sequence Difference for Insects Does Not Predict Species Boundaries

Diagnosis and assessment of species boundaries of economically important insects are often problematic because of limited morphological and/or biological characters. DNA data can help to identify and revise species. Nonoverlapping intra- and interspecific sequence divergences are often used as evidence for species. Thus, the establishment of a standardized percent nucleotide divergence to predict species boundaries would aid in cases where species status is suspect. However, given variation in nucleotide mutation rates and species concepts, association between a standard percent sequence divergence and species is questionable. This review surveys the percent DNA sequence difference found between sister-species of economically important insects, to assess whether a standard divergence associates with all taxa. Sixty-two comparisons of intra- and interspecific pairwise DNA differences were made for mitochondrial and nuclear loci spanning families of Isoptera, Phthiraptera, Hemiptera, Coleoptera, Lepidoptera, Diptera, and Hymenoptera. Intra- and interspecific sequence divergences varied widely among insects, 0.04-26.0 and 1.0-30.7%, respectively. The ranges of intra- and interspecific sequence divergences overlapped in 28 of 62 comparisons. This implies that a standardized percent sequence divergence would fail to correctly diagnose species for 45% of the cases. Common occurrence of nonmonophyly among closely related species probably explains this observation. Nonmonophyly and overlap of intra- and interspecific divergences were significantly associated. The reviewed studies suggest that a standard percent sequence divergence does not predict species boundaries among economically important insects. DNA data can help best to predict species boundaries via its inclusion in nonphenetic phylogenetic analysis and subsequent systematic expert scrutiny.

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.

100 million years of morphological conservation in bark beetles (Coleoptera: Curculionidae: Scolytinae)

Abstract Scolytine weevils (bark and ambrosia beetles) have a unique ecological significance in forest ecosystems, which equates to major effects on landscape ecology and to monetary losses. Fossilized galleries of scolytines have been reported in Late Mesozoic wood, but here we describe a well‐preserved body fossil from the Cretaceous, c. 100 Ma, preserved in amber from northern Myanmar. Moreover, the specimen is remarkably similar to Recent species of the genus Microborus, revealing stasis unexpected within scolytines and thus highlighting the antiquity of the group. Stratigraphic dating and comparison of insect palaeofaunas included in other well‐dated ambers from multiple sites support the age estimate of the Burmese amber. A minimum age for one clade of scolytines is thus established, indicating an early divergence of scolytines from other weevils in the Late Jurassic or Early Cretaceous and challenging the current perspective of weevil evolution.

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

BackgroundFungus 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.ResultsPhylogenetic 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.ConclusionOrigins 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.

Phylogeography of the bark beetle Dendroctonus mexicanus Hopkins (Coleoptera:Curculionidae: Scolytinae)

Dendroctonus mexicanus is polyphagous within the Pinus genus and has a wide geographical distribution in Mexico and Guatemala. We examined the pattern of genetic variation across the range of this species to explore its demographic history and its phylogeographic pattern. Analysis of the mtDNA sequences of 173 individuals from 25 Mexican populations allowed to us identify 53 geographically structured haplotypes. High haplotype and low nucleotide diversities and Tajimas D indicate that D. mexicanus experienced rapid population expansion during its dispersal across mountain systems within its current range. The nested clade phylogeographic analysis indicates that the phylogeographic pattern of D. mexicanus is explained by continuous dispersion among lineages from the Sierra Madre Occidental, the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt. However, we also observed isolation events among haplotypes from the Cofre de Perote/Trans-Mexican Volcanic Belt/Sierra Madre Oriental and the Trans-Mexican Volcanic Belt/Sierra Madre del Sur, which is consistent with the present conformation of mountain systems in Mexico and the emergence of geographical barriers during the Pleistocene.

Cladistic review of generic taxonomic characters in Xyleborina (Coleoptera: Curculionidae: Scolytinae)

Abstract A cladistic analysis of morphological characters of the subtribe Xyleborina (Curculionidae, Scolytinae) is presented. An examination of individual characters revealed little phylogenetic information in many characters currently used for delimiting genera. Phylogenetically stable characters were used for the evaluation of the contemporary generic concept. The following genera have been recovered as monophyletic: Cnestus, Dryocoetoides, Eccoptopterus, Xylosandrus, Schedlia, Sampsonius and Taurodemus. The following genera have been found to be polyphyletic: Amasa, Ambrosiodmus, Arixyleborus, Coptoborus, Coptodryas, Cryptoxyleborus, Cyclorhipidion, Euwallacea, Leptoxyleborus, Taphrodasus, Theoborus, Webbia, Xyleborinus and Xyleborus. The analysis permitted the resurrection of four genera: Anisandrus, Microperus, Pseudowebbia and Streptocranus. A number of new combinations at specific level are given: Anisandrus cornutus (Schaufuss, 1891), A. dispar (Fabricius, 1792), A. eggersi (Beeson, 1930), A. improbus (Sampson, 1913), A. longidens (Eggers, 1930), A. maiche Stark, 1936, A. obesus (LeConte, 1868), A. sayi Hopkins, 1915, A. apicalis (Blandford, 1894), A. hirtus (Hagedorn, 1904), Microperus myristicae (Schedl, 1939), M. eucalypticus (Schedl, 1938), M. huangi (Browne, 1983), M. intermedius (Eggers, 1923), M. kadoyamaensis (Murayama, 1934), Pseudowebbia armifer (Schedl, 1942), P. seriata Browne, 1963, P. squamatilis (Schedl, 1955), P. trepanicauda (Eggers, 1923), P. curvatus (Browne, 1986), Streptocranus bicolor Browne, 1949, S. bicuspis (Eggers, 1940), S. capucinulus (Schedl, 1942), S. forficatus (Schedl, 1957), S. fragilis Browne, 1949, S. longicauda Browne, 1960, S. longispinis Browne, 1986, S. mirabilis Schedl, 1939, S. usagaricus (Eggers, 1922), S. sexdentatus (Eggers, 1940). The characters most useful for generic‐level taxonomy of Xyleborina were identified and their states refined and illustrated. An accompanying illustrated multiple‐entry electronic key for the updated xyleborine classification has been published on‐line at www.scolytid.msu.edu.

REPEATED EVOLUTION OF CROP THEFT IN FUNGUS-FARMING AMBROSIA BEETLES

Ambrosia beetles, dominant wood degraders in the tropics, create tunnels in dead trees and employ gardens of symbiotic fungi to extract nutrients from wood. Specificity of the beetle–fungus relationship has rarely been examined, and simple vertical transmission of a specific fungal cultivar by each beetle species is often assumed in literature. We report repeated evolution of fungal crop stealing, termed mycocleptism, among ambrosia beetles. The mycocleptic species seek brood galleries of other species, and exploit their established fungal gardens by tunneling through the ambient mycelium‐laden wood. Instead of carrying their own fungal sybmbionts, mycocleptae depend on adopting the fungal assemblages of their host species, as shown by an analysis of fungal DNA from beetle galleries. The evidence for widespread horizontal exchange of fungi between beetles challenges the traditional concept of ambrosia fungi as species‐specific symbionts. Fungus stealing appears to be an evolutionarily successful strategy. It evolved independently in several beetle clades, two of which have radiated, and at least one case was accompanied by a loss of the beetles’ fungus‐transporting organs. We demonstrate this using the first robust phylogeny of one of the worlds largest group of ambrosia beetles, Xyleborini.

DNA based cladograms augment the discovery of a new Ips species from China (Coleoptera: Curculionidae: Scolytinae)

The implementation of DNA in taxonomic study is in its infancy because the association of the amount and type of nucleotide change with species boundaries has not been fully examined for most taxa. Mitochondrial cytochrome c oxidase I (COI) nucleotide data is currently the most popular molecular marker for delimiting species boundaries and a standard pair‐wise nucleotide divergence between groups of individuals has been suggested for the recognition of new species. It is unlikely that such a standard would be applicable across animal species, but the association of the amount and type of nucleotide change with species boundaries could help with the establishment of a taxon‐specific DNA taxonomy.

Morphology, Taxonomy, and Phylogenetics of Bark Beetles

This chapter summarizes the historical and contemporary taxonomy, phylogenetic relationships, morphological and molecular approaches, and information resources used in the classification of bark and ambrosia beetles (Coleoptera: Curculionidae: Scolytinae). The chapter pays special attention to the description of methods, including details on morphological and molecular characters on which the current classification is based. Illustrations and a detailed glossary of representative groups will allow users to understand the morphology of bark beetles. Relationships are illustrated by an up-to-date consensus phylogenetic tree. Diversity of the group, as well as individual tribes and genera, are exhaustively tabulated for the first time. Controversies surrounding the evolutionary origins of bark beetles and ambiguities in their morphology and classification are also discussed.