PSYCHE 
Vol. 73 June, 1966 No. 2 
PROTELYTROPTERA FROM THE UPPER PERMIAN OF 
AUSTRALIA, WITH A DISCUSSION OF THE PROTO- 
COLEOPTERA AND PARACOLEOPTERA 1 
By Jarmila Kukalova 
Charles University, Prague 
In the Tillyard collection of Upper Permian insects from Belmont, 
New South Wales (now in the British Museum of Natural History 
in London), there is a remarkable series of extinct elytrophorous in- 
sects of the order Protelytroptera. The specimens belong to four 
families, all endemic to Australia so far as known. Some of the 
genera, however, have formerly been referred to the “orders” Proto- 
coleoptera and Paracoleoptera, which were supposed to represent the 
ancestors of true Coleoptera. 
The order Protelytroptera, one of the most diversified groups of 
hemimetabolous insects of extensive geographical distribution, has 
been found so far in Permian strata of North America, Czechoslo- 
vakia, U.R.S.S. and Australia. Because of the striking similarity of 
their fore wings to those of beetles, the remarkable Australian fossils, 
Protocoleus and Permophilus , were described by Tillyard (1924) as 
primitive coleopteroids, Protocoleus being placed in a new order, 
Protocoleoptera, and Permophilus in the Coleoptera. Their phylo- 
genetic position has been repeatedly discussed in the literature. 
Although the protelytropterous character of Protocoleus was pointed 
out later by Tillyard (1931) and Carpenter (1933), the systematic 
position of Permophilus has remained uncertain and in 1953 it was 
assigned by Laurentiaux to another new order, Paracoleoptera. Nu- 
merous and well preserved specimens in the Tillyard collection in 
the British Museum have enabled me to recognize the orders Proto- 
J Much of this research was done while the author was at the Biological 
Laboratories, Harvard University, 1964, under a National Science Founda- 
tion Grant (NSF GB-2038j to Professor Carpenter, to whom I am grateful 
for help in the preparation of this paper. I am much indebted to Dr. E. I. 
White and Mr. R. Baker of the British Museum (Natural History) for the 
opportunity of studying these Australian insects. A grant-in-aid of research 
from the Society of the Sigma Xi has covered part of the cost of publication. 
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