title: The earliest evidence of holometabolan insect pupation in conifer wood.
abstract: BACKGROUND
  The pre-Jurassic record of terrestrial wood borings is poorly resolved, despite body fossil evidence of insect diversification among xylophilic clades starting in the late Paleozoic. Detailed analysis of borings in petrified wood provides direct evidence of wood utilization by invertebrate animals, which typically comprises feeding behaviors.
  
  
  RESULTS
  We describe a U-shaped boring in petrified wood from the Late Triassic Chinle Formation of southern Utah that demonstrates a strong linkage between insect ontogeny and conifer wood resources. Xylokrypta durossi new ichnogenus and ichnospecies is a large excavation in wood that is backfilled with partially digested xylem, creating a secluded chamber. The tracemaker exited the chamber by way of a small vertical shaft. This sequence of behaviors is most consistent with the entrance of a larva followed by pupal quiescence and adult emergence--hallmarks of holometabolous insect ontogeny. Among the known body fossil record of Triassic insects, cupedid beetles (Coleoptera: Archostemata) are deemed the most plausible tracemakers of Xylokrypta, based on their body size and modern xylobiotic lifestyle.
  
  
  CONCLUSIONS
  This oldest record of pupation in fossil wood provides an alternative interpretation to borings once regarded as evidence for Triassic bees. Instead Xylokrypta suggests that early archostematan beetles were leaders in exploiting wood substrates well before modern clades of xylophages arose in the late Mesozoic.
authors.affiliation: Department of Geosciences, Idaho State University, Pocatello, Idaho, United States of America. tapaleif@isu.edu
authors.forename: Leif
authors.initials: L
authors.lastname: Tapanila
authors.forename: Eric M
authors.initials: EM
authors.lastname: Roberts
mesh_terms: D000818:Animals; D005075:Biological Evolution; D007313:Insecta / Q000145:classification / Q000254:growth & development*; D008675:Metamorphosis, Biological; D011679:Pupa / Q000254:growth & development*; D064028:Tracheophyta / Q000033:anatomy & histology* / Q000254:growth & development; D014934:Wood / Q000033:anatomy & histology* / Q000254:growth & development