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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: S. praeceps LeConte, 1868 (= S. abietis Blackman, 1934; = S. opacus Blackman, 1934), S. reflexus Blackman, 1934 (= S. virgatus Bright, 1972; = S. wickhami Blackman, 1934). Two species were reinstated: S. fiskei Blackman, 1934 and S. silvaticus Bright, 1972. A diagnosis, description, distribution, host records and images were provided for each species and a key is presented to all species.
Natural history 4
Sexual dimorphism 8
Systematics 9
Taxonomic history 9
Materials 10
Morphology 10
Taxonomic characters 12
Taxa, DNA sequences and alignment 18
Methods 27
Species concept 27
Morphology 28
Molecular 28
Distribution mapping 31
Terminology 31
Phylogenetic results 35
Morphology 35
Mitochondrial COI 35
Combined analyses 35
Taxonomic treatment 38
A checklist of the Nearctic Scolytus species 38
Scolytus Geoffroy, 1762 39
Scolytus mali (Bechstein, 1805) 40
Scolytus multistriatus (Marsham, 1802) 45
Scolytus rugulosus (Müller, 1818) 50
Scolytus schevyrewi Semenov, 1902 55
Native Hardwood clade 60
Scolytus fagi Walsh, 1867 60
Scolytus muticus Say, 1824 64
Scolytus quadrispinosus Say, 1824 69
Conifer clade 74
Scolytus aztecus Wood, 1967 74
Scolytus dentatus Bright, 1964 80
Scolytus fiskei Blackman, 1934, valid sp. 83
Scolytus hermosus Wood, 1968 90
Scolytus laricis Blackman, 1934 94
Scolytus monticolae (Swaine, 1917) 99
Scolytus mundus Wood, 1968 104
Scolytus obelus Wood, 1962 107
Scolytus oregoni Blackman, 1934 111
Scolytus piceae (Swaine, 1910) 114
Scolytus praeceps LeConte, 1876 120
Scolytus reflexus Blackman, 1934 127
Scolytus robustus Blackman, 1934 134
Scolytus silvaticus Bright, 1972 valid sp. 138
Scolytus subscaber LeConte, 1876 141
Scolytus tsugae (Swaine, 1917) 144
Scolytus unispinosus LeConte, 1876 149
Scolytus ventralis LeConte, 1868 156
Key to the Nearctic adults of Scolytus 161
Discussion 171
Acknowledgements 172
References 173