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      A taxonomic monograph of Nearctic Scolytus Geoffroy (Coleoptera, Curculionidae, Scolytinae)

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          Abstract

          Abstract

          The Nearctic bark beetle genus Scolytus Geoffroy was revised based in part on a molecular and morphological phylogeny. Monophyly of the native species was tested using mitochondrial (COI) and nuclear (28S, CAD, ArgK) genes and 43 morphological characters in parsimony and Bayesian phylogenetic analyses. Parsimony analyses of molecular and combined datasets provided mixed results while Bayesian analysis recovered most nodes with posterior probabilities >90%. Native hardwood- and conifer-feeding Scolytus species were recovered as paraphyletic. Native Nearctic species were recovered as paraphyletic with hardwood-feeding species sister to Palearctic hardwood-feeding species rather than to native conifer-feeding species. The Nearctic conifer-feeding species were monophyletic. Twenty-five species were recognized. Four new synonyms were discovered: Scolytus praeceps LeConte, 1868 (= Scolytus abietis Blackman, 1934; = Scolytus opacus Blackman, 1934), Scolytus reflexus Blackman, 1934 (= Scolytus virgatus Bright, 1972; = Scolytus wickhami Blackman, 1934). Two species were reinstated: Scolytus fiskei Blackman, 1934 and Scolytus silvaticus Bright, 1972. A diagnosis, description, distribution, host records and images were provided for each species and a key is presented to all species.

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          The evolution of agriculture in beetles (Curculionidae: Scolytinae and Platypodinae).

          Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi--known as ambrosia fungi--that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free-living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle-fungus associations, enabling them to use the biomass-rich resource that trees represent and set the stage for at least one origin of eusociality.
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            How do insect nuclear and mitochondrial gene substitution patterns differ? Insights from Bayesian analyses of combined datasets.

            We analyzed 12 combined mitochondrial and nuclear gene datasets in seven orders of insects using both equal weights parsimony (to evaluate phylogenetic utility) and Bayesian methods (to investigate substitution patterns). For the Bayesian analyses we used relatively complex models (e.g., general time reversible models with rate variation) that allowed us to quantitatively compare relative rates among genes and codon positions, patterns of rate variation among genes, and substitution patterns within genes. Our analyses indicate that nuclear and mitochondrial genes differ in a number of important ways, some of which are correlated with phylogenetic utility. First and most obviously, nuclear genes generally evolve more slowly than mitochondrial genes (except in one case), making them better markers for deep divergences. Second, nuclear genes showed universally high values of CI and (generally) contribute more to overall tree resolution than mitochondrial genes (as measured by partitioned Bremer support). Third, nuclear genes show more homogeneous patterns of among-site rate variation (higher values of alpha than mitochondrial genes). Finally, nuclear genes show more symmetrical transformation rate matrices than mitochondrial genes. The combination of low values of alpha and highly asymmetrical transformation rate matrices may explain the overall poor performance of mitochondrial genes when compared to nuclear genes in the same analysis. Our analyses indicate that some parameters are highly correlated. For example, A/T bias was positively and significantly associated with relative rate and CI was positively and significantly associated with alpha (the shape of the gamma distribution). These results provide important insights into the substitution patterns that might characterized high quality genes for phylogenetic analysis: high values of alpha, unbiased base composition, and symmetrical transformation rate matrices. We argue that insect molecular systematists should increasingly focus on nuclear rather than mitochondrial gene datasets because nuclear genes do not suffer from the same substitutional biases that characterize mitochondrial genes.
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              Systema entomologiae : sistens insectorvm classes, ordines, genera, species, adiectis synonymis, locis, descriptionibvs, observationibvs / Io. Christ. Fabricii.

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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2014
                29 October 2014
                : 450
                : 1-182
                Affiliations
                [1 ]Department of Entomology, Michigan State University, Natural Science Building, room 243, 288 Farm Lane, East Lansing, MI 48824, United States of America
                Author notes
                Corresponding author: Sarah M. Smith ( camptocerus@ 123456gmail.com )

                Academic editor: M. Alonso-Zarazaga

                Article
                10.3897/zookeys.450.7452
                4233402
                11a6e786-5c26-4c48-9f8d-88c625352746
                Sarah M. Smith, Anthony I. Cognato

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 7 March 2014
                : 13 August 2014
                Categories
                Monograph

                Animal science & Zoology
                scolytidae,bark beetle,fir engraver,banded elm bark beetle,lesser elm bark beetle,douglas-fir engraver,hickory bark beetle,shot hole borer,large shot hole borer,hackberry engraver,taxonomy,revision,animalia,coleoptera,curculionidae

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