90 
Psyche 
[June 
coleoptera and Paracoleoptera as synonyms of the Protelytroptera 
and to describe additional members of Australian Protelytroptera . 2 
Order Protelytroptera Tilly ard 
Family Protocoleidae Tillyard, 1924 
Protocoleidae Tillyard, 1924, Proc. Linn. Soc. N.S.W. 49: (4) :434. Lameere, 
1932, Soc. ent. Br., Liv. Cent.: 596; Forbes, 1928, Psyche, 3 5 :33 ; Tillyard’ 
1931, Amer. Journ. Sci. 21:234; Peyerimhoff, 1934, Bull. Soc. ent. Fr. 39:39; 
Richter, 1936, Rev. d’ent. URSS, 26:31; Lameere, 1938, Bull. Ann. Soc. 
ent. Belg. 78:355; Jeannel, 1949, Traite de Zool. 9:63; Laurentiaux, 1953, 
Traite de Paleont. 3:475; Carpenter, 1954, Classification of Insects, p. 789 ; 
Rohdendorf, 1962, Osnovy paleont., p. 268. 
Diagnosis: The following diagnosis of the family is based on the 
description and photograph of Protocoleus mitchelli Tillyard, as well 
as on a study of the topotypical material of Protocoleidae in the 
British Museum. 
Fore wings: large, tegminous, little sclerotized, only slightly con- 
vex; sutural margin bordering the whole posterior margin; wing 
surface covered with dense granulation and larger flat tubercles, ap- 
parently on both dorsal and ventral surfaces ; a cluster of setae present 
in the subcostal area; costal expansion large, projecting, rounded; 
anterior margin strongly convex; venation richly branched, especially 
the radius; Sc long, dividing into several parallel branches; R send- 
ing off a series of branches anteriorly; Rs short or missing; M and 
CuA variable in form; CuP slightly concave or flat, often branched; 
anal veins directed anteriorly, 3-5 in number; cross veins numerous, 
regular or irregular, sometimes connected by anastomoses. 
Relationship. In 1924 Tillyard described a remarkable fossil fore 
wing as Protocoleus mitchelli and referred it to his new order Proto- 
coleoptera. In his opinion this order was intermediate between the 
Carboniferous Protoblattoidea and the Coleoptera. Subsequently, 
much discussion has centered on the phylogenetic and systematic posi- 
tion of Protocoleus. Many authors (e.g., Lameere, 1926, Forbes, 
1928, Peyerimhoff, 1933, Carpenter, 1933, Richter, 1935, Jeannel, 
1949) doubted the relationship of Protocoleus to the true Coleoptera. 
Tillyard himself in 1931 (p. 264) stated that the Protocoleidae could 
not have been the real ancestors of the Coleoptera but were more 
probably an archaic remnant of the older Protelytroptera. Lauren- 
tiaux (1953, p. 475) put this genus in the distinct order Proto- 
coleoptera but agreed with its proximity to Dermaptera and 
Tor a general account of the Protelytroptera, see Carpenter and Kukalova, 
1964, and Kukalova, 1965. 
