A Dendroctonus bark engraving (Coleoptera: Scolytidae) from a middle Eocene Larix (Coniferales: Pinaceae):: Early or delayed colonization?

An engraving made by a scolytid bark beetle, assigned to the genus Dendroetonus of the tribe Tomieini, has been identified on a mummified, middle Eocene (45 Ma) specimen of Larix altoborealis wood from the Canadian High Arctic. Larix altoborealis is the earliest known species of Larix, a distinctive lineage of pinaceous conifers that is taxonomically identifiable by the middle Eocene and achieved a broad continental distribution in northern North America and Eurasia during the late Cenozoic. Dendroctonus currently consists of three highly host-specific lineages that have pinaceous hosts: a basal monospecific clade on Pinoideae (Pinus) and two sister clades that consist of a speciose clade associated exclusively with Pinoideae and six species that breed overwhelmingly in Piceoideae (Picea) and Laricoideae (Pseudotsuga and Larix). The middle Eocene engraving in L. altoborealis represents an early member of Dendroctonus that is ancestral to other congeneric species that colonized a short-bracted species of Larix. This fossil occurrence, buttressed by recent data on the phylogeny of Pinaceae subfamilies and Dendroetonus species, indicates that there phylogenetically congruent colonization by these bark-beetle lineages of a Pinoideae + (Piceoideae + Laricoideae) host-plant sequence. Based on all available evidence, an hypothesis of a geochronologically early invasion during the Early Cretaceous is supported over an alternative view of late Cenozoic cladogenesis by bark beetles onto the Pinaceae. These data also suggest that host-plant chemistry may be an effective species barrier to colonization by some bark-beetle taxa over geologically long time scales.