e-journal
Evolution of insect wings and development – new details from Palaeozoic nymphs
The nymphal stages of Palaeozoic insects differ significantly in morphology from those of their modern counterparts.
Morphological details for some previously reported species have recently been called into question. Palaeozoic insect
nymphs are important, however – their study could provide key insights into the evolution of wings, and complete
metamorphosis. Here we review past work on these topics and juvenile insects in the fossil record, and then present both
novel and previously described nymphs, documented using new imaging methods. Our results demonstrate that some
Carboniferous nymphs – those of Palaeodictyopteroidea – possessed movable wing pads and appear to have been able
to perform simple flapping flight. It remains unclear whether this feature is ancestral for Pterygota or an autapomorphy
of Palaeodictyopteroidea. Further characters of nymphal development which were probably in the ground pattern of
Pterygota can be reconstructed. Wing development was very gradual (archimetaboly). Wing pads did not protrude
from the tergum postero-laterally as in most modern nymphs, but laterally, and had well-developed venation. The
modern orientation of wing pads and the delay of wing development into later developmental stages (condensation)
appears to have evolved several times independently within Pterygota: in Ephemeroptera, Odonatoptera, Eumetabola,
and probably several times within Polyneoptera. Selective pressure appears to have favoured a more pronounced
metamorphosis between the last nymphal and adult stage, ultimately reducing exploitation competition between the
two. We caution, however, that the results presented herein remain preliminary, and the reconstructed evolutionary
scenario contains gaps and uncertainties. Additional comparative data need to be collected. The present study is thus seen as a starting point for this enterprise.
Key words: insect development, fossils, nymphs, ontogeny, palaeo-evo-devo, palaeoentomology.
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