Jarmila Kukalová.
Protelytroptera from the Upper Permian of Australia, with a Discussion of the Protocoleoptera and Paracoleoptera.
Psyche 73:89-111, 1966.

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PSYCHE
Vol. 73 June, 1966 No. 2
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PROTELYTROPTERA FROM THE UPPER PERMIAN OF AUSTRALIA, WITH A DISCUSSION OF THE PROTO- COLEOPTERA AND PARACOLEOPTERA1
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 Permo$hilus, were described by Tillyard (1924) as primitive coleopteroids, Protocoleus being placed in a new order, Protocoleoptera, and Permophihis 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- '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-2038; 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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90 Psyche [June
~oleo~ptera and Paracoleoptera as synonyms of the Protelytroptera and to describe additional members of Australian Pr~telytroptera.~ Family Protocoleidae Tillyard, I 924
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. 78 555 ; Jeannel, 1949, Trait6 de 2001. 9 :63 ; Laurentiaux, 1953, Trait6 dc 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 mitchelll 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 loiig, 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 01- 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 mitchell'i and referred it to his new order Proto- coleoptera. In his opinion this order was intermediate between the Carboniferous Protoblattoidea and the Coleo~tera. Subsequently, much discussion has centered on the ph~logenetic 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 01 Protocolezu 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. L'auren- tiaux (1953, p. 475) put this genus in the distinct order Proto- coleoptei-a but agreed with its proximity to Dermaptera and Tor a general account of the Protelytroptera, see Carpenter and Kukalovi, 1964, and Kukalovi, 1965.




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19661 Kukdovd - Protelytroptera 91
Protelytroptera. Carpenter (1954, p. 798) referred the family to the Protelytroptera, which he considered to be a highly specialized order, resembling the Coleoptera but actually related to the Blat- todea. Rohdendorf ( I 962) referred the family to the Coleopteroidea, incertae sedis.
In the Tillyard collection at the British Museum there are 7 com- plete fore wings of the family Protocoleidae well preserved enough to confirm the correctness of Carpenter's conclusion (1954). The specimens show the prominent, typical costal expansion (not pre-
served in the holotype of P. mitchelli), a well developed sutural mar- gin, a distinct patch of setae and, in spite of the unusual richness of branches, a protelytropterous pattern of venation. The family Protocoleidae includes the largest forms among Pro- telytroptera and is well removed from all other families, excepting the Permophilidae, which also have the wing surface covered with both granulation and tubercles, a similar outline of the wing and a broad subcostal area.
Two other species of elytrophorous insects were described by Till- yard from the Permian of New South Wales: Permofulgor bel- montens'is Tillyard ( 19 I 7) and Permofulgor indistinctus Tillyard (1922). The family Permofulgoridae was established for the genus Permofulgor by Tillyard in 1917 and was placed in the order Hemip- tera. Subsequently (1926, p. 186) Tillyard became convinced that these fossils were related to Protocoleus and assigned them to the Protocoleoptera. They are presumably protelytropterous, also, but because of poor preservation and the fragmentary nature of these fossils, generic and family diagnoses cannot be made. Since the venation of the Protocoleidae is both complicated and indistinct, drawings are unusually difficult to make. Fortunately, venational details are very variable and of little use at the specific level.
Geological occurrence of the family: Upper Permian of Australia (N.S.W.)
Genera included: Protocoleus Tillyard, 1924; Phyllelytron, new genus ; hstrelytron, new genus.
Genus Protocoleus Tillyard
Proiocoleus Tillyard, 1924, Proc. Linn. Soc., N.S.W. 49 (4) :434; Tillyard, 1931, Amer. Journ. Sci. 21 234.
Diagnosis. Fore wing: tegminous, darkly pigmented; wing surface covered by flat, regularly arranged tubercles (not occurring on veins)



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and by fine granulation; costal expansion unknown; wings relatively broad and short, apical part prolonged and narrow; anterior margin strongly convex, posterior margin slightly concave apically; branches of main veins nearly parallel, distinct; Sc sending off several parallel branches, almost leaching apex; R sending off a series of long branches towards Sc; Rs hardly distinguishable from the other branches of R; M and CuA with a variable number of irregular branches; CUP long, branched, slightly concave; anal veins about 5 in number, some directed anteriorly; cross veins numerous, regular. Relationship. Protocolens Tillyard, I 924, differs from the related new genus Phyllelytron in the more distinct venation, in the more regularly and parallel-arranged veins and cross veins, and in the regularly arranged tubercles, distinct on the whole wing surface. From Austrelytron, n. gen., it differs in the more richly branched main veins, the presence of many cross veins and absence of regular series of pointed tubercles on the veins.
Stratigraphic occurrence : Upper Permian of Australia (N.S.W.) . Type-species : Protocoleus mitchelli Tillyard, 1924 Pro tocoleus mitchelli Tillyard
Protocoleus mitchelli Tillyard, 1924, Proc. Linn. Soc., N.S.W. 49(4) :432; pi. 46, fig. 3.
Fore wing: about 27-29 mm. long and 7-9.3 mm. broad, sutural margin broader in the proximal half, narrowing abruptly in the distal half; tubercles densely arranged, often bordering veins in regular rows; fore wing very narrow in the apical third; anterior margin very convex, in the distal third slightly concave; branches of R dis- tinct, very oblique and regular ; cross veins mostly straight. Discussion. Since Tillyard had at his disposal only the reverse of Protocoleus for study, he considered the surface of the elytron to be covered by pits. Actually, the tubercles and interspaces between them are so much alike that the reverse and obverse sculpturing does not show much difference. The fine granulation, on the other hand, ap- pears in the reverse half of the fossil as small, dense pits. The granulation, which is extraordinarily fine in Protocoleus, was not mentioned in Tillyard's original description. It is very probable that both tubercles and granulation were present on both dorsal and ventral surfaces of the tegmina,
Holotype: specimen (reverse), figured by Tillyard ( I 924) on plate 46, fig. 3 ; collected in Upper Permian beds of Belmont, N.S.W. (Not seen).




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Text-figure 1. Phyllelytron folium n. sp. Drawing of holotype, No. In 45364 Brit. Mus. ; fore wing, Upper Permian, Australia (original). Let- tering: Sc, subcosta; Rl, radius; Rs, radial sector; M, media; CuA, an- terior cubitus ; CUP, posterior cubitus; Al, first anal vein ; An, anal area. Text-fignre 2. Phyllelytron 9etalbn n. sp. Drawing of holotype, No. In 45703, Brit. Mus. ; fore wing, Upper Permian, Australia (original). Text-figure 3. Phyllelytron qranulatum n. sp. Drawing of holotype, NO, In 38115, Brit. Mus.; fore wing, Upper Permian, Australia (original).



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94 Psyche [June
Genus Phyllelytron, new genus
Fore wing : tegminous) darkIy pigmented ; sutural margin narrow ; tubercles distinct in the anal area or in the whole posterior part of the wing, flat; granulation dense, distinct on the whole wing surface; costal expansion of circular shape, strongly projecting; apex directed posteriorly, pointed, apical part only little prolonged ; anterior niar- gin convex; veins not running parallel, branches of R, h!l and CuA usually indistinct; Sc sending ofi 2 or more long parallel branches, with numerous veinlets ; R sendkg off anteriorly a series of irregular, slightly oblique, weak branches; M and CuA of various forms, branched ; CUP long, slightly concave, usually branched ; anal veins broad, 4-5 in number, some of them directed anteriorly; cross veins nunlesous, irregular.
Relationship. PhyIZeZytron, n. gen., differs from ProtocoIeus Till- yard in the less prolonged apex, more irregular branches of veins (mzinly R), less distinct veins) less numerous tubercles restricted mainly to the posterior half of the wing, not regularly arranged; granulztion more coarse and cross veins more irregular. PhylZ~~Zytron, n. gen.) differs from Austrelytron, n. gen., in the moi-e richly branched keins and more mmerous cross veins, smaller number of anal veins and flat tubercles, which are absent on the veins. Stratigraphic occurrence : Upper Permian of Australia (N.S.W. ) Type-species : Phy ZZeZytron f olium n. sp. Phyllelytron folium, 11. sp.
Text-figure I
Fore wing: 21-24 nlm. long, 6.2-7.8 mm. broad; sutural margin relatively narrow; tubercles distinct in the anal area, in the other parts of the wing almost n~issing; granulation fine) costa1 expansion markedly projecting, circular ; anterior margin strengthened ; pos- terior margin slightly cmcave) with small concavity in the anal part; branches of R indistinct) irregular, connected with few cross veins; Relationship. Phyllelytrwz foliunz, n. spa) differs from P, gvtdon, n. sp.) in the more regular) less numerous cross veins, in the absence of Jark color spots in the proximal half of the wing and in the equally dark-pigmented veins.
In comparison with P. grunuIutz~~~z,
n. sp., it h2s a more projecting costa1 expansion) smaller and less coarse granulatior~s, more distinct tubercles and no dark spots. From P. nzelinzm, n. sp., it differs in its shorter, broader and probably thinner tegnlen) narrower sutural margin and more psojecting costal expansion.




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Holotype: Na. In 45364 (reverse). Specimens No. In 45529 and In 4.5504 are appzsentlv the same species. British Museum (N.H.), London. Collected in Upper Permian, Belmont, Australia (N.S.1V.). Phyllelytron petalon, n. sp.
Text-figure 2
Fore wing: 19 mm. long) 6.5 broad ; sutural margin considerably narrow; tubercles we11 developed on the whole posterior half of the wing; granulatiori medium coarse; costal expansion projecting) cis- culm-; branches of R slightly oblique, irregular, connected with many cross veins; cross veins numerous, very irregular; veins and csoss veins not pigmented, interspaces between them darkly pigmented before all in the distal half of the wing; darker spot before the mid- dle of the wing, along the basal part of the anterior margin and round the base.
Relationship. PhyZZeZytron fietalon, n. sp., is related to P. qrunu- lafiunz, n. ,spa, by darkly colored tegmina with unpigmented veins. It differs from this species in its finer granulation) distinct tubercles and relatively large, different pattern of colored spots and irregular cross veins.
Holotype: No. In 45703 (obverse). Specimens No. In 45528 and 45734 are probably this species. British Museum (N.H.)) Lon- don. Collected in Upper Permian, Belmont) N.S.W. Phyllelytron granulatum, n. sp.
Text-figure 3
Fose wing: 15.2 mm. long and 5.3 rnm. broad; sutural margin narrow ; tubei-cles indistinct ; granulation coarse ; costal expansion projecting; anterior margin strengthened) pigmented in the proximal half; branches of R indistinct, irregular) connected with cross veins; cross veins not very numerous, irregular; veins not pigmented, three dark colored spots in the proximal half of the wing. Relationship: PhyZZelytron g~~nulatunz, n. sp., differs from all species of the same gem{$ in smallest dimensions) coarsest granulation a d indistinct tubercles.
Holotype : No, In 381 13 (revei-se). Specimens No. In 455 I 3 and In 45519 are the same species. British Museum (N.H.)) London. Collected in Upper Permian) Belmont) N.S.W. Phyllelytron melinum, n. sp.
Text-figure 4
Fore wing: about 28 mm, long) 6.8 mm. broad, uniformly dark- colored; sut~lral margin broad; tubercles well developed only in the



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96 Psyche
Text-figure 4.
Phylielytron melinum n, sp. Drawing of holotyye, No. In 45510, Brit. hius.; fore wing, Upper Permian, Australia (original). anal area; granulation fine; costal expansion not very projecting; branches of R indistinct, irregular, connected with few cross veins; RI, A1 and CuA very indistinct; cross veins faintly visible, few, mostly regular.
Relationship. PhyZZeZytron melinurn, n. sp., differs from all species of the genus in iargest, relatively narrowest and apparently thickest tegmina with broadest sutural margin and very indistinct RI, M and CuA.
Holotype :
No. In 455 10 (obverse). British Museum (N.H.) , Lorldon. Collected in Upper Permian, Belmont, N.S.W. Genus Austrelytron, new genus
Fore wing: tegminous, relatively small ; sutural margin narrow ; wing surface covered by dense granulation and isolated, pointed tubercles, which also form rows on the veins; costal expansion not very pro-jecting; wings broadest before the middle of the wing) nar- rowing quickly towards the apex; anterior margin convex, posterior margin slightly concave; main veins with few branches; Sc with t m 7 0 or more branches parallel to each other, terminating well before apey; R sending off several irregular branches towards Sc; Rq dis- tinct) originating after the middle of the wing; CUP long) concave; about three unbranched anal veins; cross veins few, simple. Text-figure 5.
Ausfrelyt~on tillyardi n. sp. Drawing of holotype, No. In 45525, Brit. Mus.; fore wing, Upper Permian, Australia (original). Text-figure 6. Pcr~nophilus hirtus n. sp. Drawing of holotype, No. In 4.6014, Brit. Mus. ; fore wing, Upper Permian, (original). Text-figure 7,
Permophilus caflulus n. sp. Drawing of holotype, No, In 45518, Brit. Mus.; fore wing, Upper Permian, Australia (original). Text-figure 8. Elytrathrix hirsuta n. sp. Drawing of holotype, No. In 45503, Brit. Mus.; fore wing, Upper Permian, Australia (original).



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9s Psyche [June
Relationship. Amtrelytron, n. gen.) differs from the genus Proto- coleus and PhylZdytron, n. gen., by having fewer branches on veins, less numerous cross veins and by the nature of the tubercles) which are not flat and numercus, but pointed and isolated, and also ar- ranged in rows or1 the veins.
Stsatigraphic occurrence : Upper Permian of Austsalia (N.S.W.) . Type-species : A3ustreZytron tillyardi n. sp. Austrelytron tillyardi, n. sp,
Text-figure 5
Fore wing : I I mm. long, 4 mm. broad ; sutural margin narrow ; tubercles pointed, forming regulas rows on the veins; the ax-eas be- tween the veins covered by dense granulation and very isolated) ir- ~egulai-ly distsibuted tubercles; costal expansion projecting only a little; branches of R very irregular) distinct, connected with few ci-oss veins; all main veins distinct; R straight, RI directed anteriorly towards costa1 margin well before apex; Rs well developed; M branched, fused ior a long distance with the simple CuA; CUP long? straight) concave; anal veins simple, of irregular form. Holotype: Nc. In 45525 (obverse). British Museum (N.H.) ) London. Collected in Upper Permian) Belmont, N.S.W. Fanlily Permophilidae Tillyard, 1924
Permophilidae Tillyard, 1924, Proc. Linn. SOC., N.S.W. 49 1434 (Coleoptera) ; Laurentiaux, 1953, Trait6 de pal6ont. 3 :475 (Paracoleoptera) ; Carpenter? 1954, Classification of Insects, p. 824 (Coleoptera) . Diagnosis. Fore wing: tegminous, only slightly convex; sutural margin well developed; wing surface covered on both dorsal and ventral surfaces by dense granulation and isolated, more or less dis- tinct tubercles, sometimes more frequent in the basal part; setae present in a distinct patch ; costal expansion projecting, circular; wings narrowed abruptly before the pointed apex; venation much reduced) restricted to the basal part of the wing? in the distal part completely missing.
Relationship. Tillyard ( 1924) established the new family Permo- philidae, which he considered to be coleopteroid, ancestral to recent Hydrophilidae. According to his description, the type-species, Permo- $hih fiinconzb~i Tillyard, 1924, is large in size (21 mm.) with pointed apex, without definite longitudinal striae, but with traces of delicate, branching venation hese and there, with the surface fur- nished in several places with very weakly formed, flattened tubercles.



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In the 'Tillyard collection of the British Museum (N.H.) in Lon- don, I found 4 specimens, corresponding to this description. One of them (No. In 45518) is designated by the label, attached to the fossil and in Tillyard's hand-writing, as 1'Pernzophi2u~?'J. For that reason, there is little doubt that the fossils belong to the family Per- rnophilidae. They are clearly protelytropterous, showing the costal expansion (interpreted by Tillyard as an allula), distinct sutural margin, the setae in patches, and, in spite of its reduction, a protely- tropterous patterfi of the venation. Laurentiaux, it should be noted (1953, p. 475), referred the Permophilidae to a new order Para- coleoptera, which is not distinct from the Protelytroptera. The wing surlace of Permophilidae (dense granulation and tu- bercles), as well as the form ~f the wings, resembles that in the iamily Protocoleidae.
It is not impossible that they might have originated from a common ancestor.
Geographical cccurrence of family:
Upper Permian of Australia
(N.S.W.).
Genera includcd : Per7tzo~hilus Tillyard, 1924 ; EZytrathrixJ new genus.
Genus PermophiZus Tillyard
Pcrmophilus Tillyard, 1924, Proc. Linn. Soc., N.S.W. 49 :434 ; Jeannel, 1949, Trait6 de zoo]. 9~64; Laurentiaux, 1953, Trait6 de palkont. 3~476. Fore wing: tegmhoss, slightly convex, darkly pigmented ; sutural margin relatively narrow; wing surface covered by dense granulation and isolated: indistinct tubercles; costal expansion projecting; apical part narrowed and prolonged; main veins strongly developed in the basal third! but absent compIete1y in the distal part of the wing. Relationship. Not hning at my disposal the type for ,studyy I am not able to decide definitely the generic position of Specimens No. In 46014 and In 45518. According to the original diagnosis given by Tillyard) they differ from the holotype of P. pincombei only in the smaller size. For this I-eason, I refer them to the same genus, but they might also be considered as representing a separate genus. Speci- men No. In 455 I 7 (fragment corresponds to about the size given for the type of Permophilus, but it is too poor for any conclusion. Specimen No. In 45503, which I refer to a new genus EZytrathix, has very distinct, large tubercles concentrated at the basal part of the fore wing, which wdd presumably have been mentioned in Till- yard's description of PermophiZus pinco,mbei if they had been present. Stratigraphic occurrence: Upper Permian of A~istralia (N.S.lV.) Type-species : PermophiZus pincombti Tillyard) I 924



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Psyche
Permophilus hirtus, n. sp.
Text-figure 6
Fore wing: 6.8 mm. long and 2.2-2.4 mm. broad; granulation dense and small, covering the whole wing surface, tubercles very flat and sparse, indistinct; costal expansion unknown; apical part of wing nai-row, prolonged, the apex placed in about the longitudinal axis of the wing; anrerior margin convex, slightly concave before the apex, strengthened in the basal half; posterior margin slightly con- cave; Sc well devehped, narrowing distally to the anterior margin, reaching almost the middle of the wing length; subcostal area very broad ; R, M + CuA, Al and A2 strong veins in the basal third, but completely reduced in the more distal part of the wing; CUP con- cave, indistinct.
Relationship. PennophiZus hirtus, n. sp., differs from Permo~hilus capz~lu~, n. sp., in its smaller dimensions, more abruptly narrowing wing distally, indistinct, shortened CUP and probably more sclerotized wings and veins. From PermophiZus pincornbei Tillyard it differs by its smaller dimensions.
Remarks. In the anal area of the holotype (No. In 46014, ob- verse) a small piece of the dorsal fore wing membrane is missing so that the sculpturing of the inner side of the ventral fore wing mem- brane is visible. It is covered by punctations, which correspond to the small granulation on the outer surface of the ventral membrane. This fortunate breaking zway of the wing membrane shows that both sides of the tegmina of Permophilidae were covered by projecting sculpturing, even the ventral surface. The only punctate sculpturing on the wing surface is the small trace at insertion of setae, arranged in patches, as is to be expected in the case of good preservation in the subcostal area and along the anterior margin in the basal part of the wing.
Holotype: No. In 46014 (obverse), British Museum (N.H.) , London. Specimen No. In 45805 is the same species. Collected in
Upper Permian, Belmont, N.S.W.
Permophilus capulus, n. sp.
Text-figure 7
Fore wing: about 15 mm. long and 3.9 mm. broad, little scle- rotized ; granulation small, dense on the whole wing surface, tubercles flat and indistinct ; costal expansion unknown ; anterior margin slightly convex, strengthened only in the basal part; Sc, R, M+CuA and A1 strong, ending in the basal third; CUP concave, distinct, running its whole course; A2 and A3 also complete.



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Text-figure 9. Stenelytron enervatum n, sp. Drawing of holotype, No. In 45958 (obverse) and In 45515 (reverse), Brit. Mus.; fore wing, Upper Permian) Australia (original).
Text-figure 10. Xenclytron ligula n. sp. Drawing of holotype, No. In