J. Kukalová-Peck.
Wing-folding in the Paleozoic Insect Order Diaphanopterodea (Paleoptera), with a Description of New Representatives of the Family Elmoidae.
Psyche 81:315-333, 1974.

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WING-FOLDING 'IN THE PALEOZOIC INSECT ORDER DIAPHANOPTERODEA (PALEOPTERA), WITH A
DESCRIPTION OF NEW REPRESENTATIVES
OF THE FAMILY ELMOIDAE
BY JARMILA KUKALOVL- PECK^
Department of Geology, Carleton University, Ottawa, Ontario, Canada
The order Diaphanopterodea is closely related to two other Paleo- zoic paleopterous orders, the Palaeodictyoptera and the Megasecoptera, all three having elongate, haustellate mouth-parts. In the history of the study of paleopterous insects, the Diaphanopterodea represent the most intriguing group. The order combines the basic character- istics of the Paleoptera, fluted wings and very long, multisegrnented cerci, with the ability to fold the wings backwards over the abdomen, which is the principal character of the Neoptera. Since that resting position of the wings indicates a major development in the evolution of insects, as well as major achievement in adaptation to more varied environment, its occurrence within the Paleoptera is of great interest. At present there is general agreement among students of insect evolution that wing-folding ability arose at least twice in insects - in the Neoptera and in the Diaphanopterodea among the Paleoptera (Rohdendorf 1962, Carpenter 1963, Sharov 1971). It has 'been assumed that the wing-folding mechanism in the Diaphanopterodea was probably different from that in the Neoptera, but the structure of the axillary region of the diaphanopteran wing has not actually been known. While examining all known specimens of 'Diaphanop- terodea from the Lower Permian deposits of 'Czechoslovakia, Kansas and Oklahoma, I succeeded in finding a dozen wings, belonging to the families Elmoidae, Martynoviidae and Asthenohyrnenidae, with the bases of the wings at least partially preserved. The present paper is concerned only with the family Elmoidae; the two remaining 'This research has been supported in part by the President's Research Grant (Canadian National Research Council) from Carleton University, and in part by NSF Grant G.B. 39720, F. M. Carpenter, Harvard Uni-
versity, Principal Investigator.
I am
deeply indebted to Professor F. M. Carpenter for placing at my disposal the Diaphanopterodea in the Museum of Comparative Zoology which he had collected in Kansas and Oklahoma, and for his critical review of this manuscript.
Manuscript received by the editor June 20, 1974.



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families and the wing base morphology of the Palaeodictyoptera and Megasecoptera will be discussed elsewhere. The new material includes six new genera and ten new species of Elmoidae from the Lower Permian beds near Obora in Moravia, Czechoslovakia. The base of the wing is well preserved in one specimen only, Permodiapha carpenteri n.sp. ( No. 8/ I 974) , and incompletely preserved in seven other specimens. Since no definitive studies of the wing bases of the extant Paleop- tera [Odonata and Ephemeroptera] have been made, some difference of opinion exists about the interpretation of the basal sclerites. The interpretation of the diaphanopteran structures used in this paper should be considered only as tentative. The terminology employed is derived from the account of the wing base of Odonata by Tannert (1958), since the fore wing base in the Ephemeroptera has never been described in functional terms. For further comparison, the wing base structures of the related family Martynoviidae (un- described material) and the related order Palaeodictyoptera (Kuka- lovi 1960, 1969, 1970) have been used.
Family Elmoidae
Elmoidae Tillyard 1937: 82; Carpenter 1943: 56; Carpenter 1947: 27; Rohdendorf 1962 : 69 ; Carpenter 1963 : 249. Parelmoidae Rohidendorf 1962: 71.
The family Elmoidae was originally assigned by Tillyard to the paleopterous order Megasecoptera because of the similarity in the venation pattern. This point of view was followed by Carpenter in his emendation of Elmoidae in 1943 and 1947, although he sub- sequently ( 1954) separated the Elmoidae, along with the Diapha- nopteridae and related families, into a distinct suborder (Parame- gasecoptera), on the basis of the folded, resting position of the wings. Later, Rohdendorf ( I 962) and Carpenter ( I 963 ) independ- ently reached the conclusion that these families had to be referred to the separate order Diaphanopterodea, established by Handlirsch ( 1919). A detailed evaluation of the phylogenetic position of Elmoi- dae and Diaphanopteridae within the order, and of Diaphanopterodea within the Paleoptera has been given by Carpenter (1963). Until now, the family has been known by only three genera, all from the Lower Permian of Kansas and Oklahoma - Elmoa, Parelmoa, and Pseudelmoa, represented by five species. The new fossils from Moravia contribute additional and varied morphological features, which broaden the family characteristics formerly used. In the Moravian material thickened cuticular spots are sometimes pres-



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Kukalova-Peck - Diaphanopterodea
Fig. 1. Elmodiapha ovata nsp., specimen 1/1974, fore wing. 'Holotype. Fig. 2. Elmodiapha ovata nsp., specimen 2/1974, fore wing. Fig. 3. Pro- todiapha maculifera n.sp., specimen 3/1974, hind wing. Holotype. Fig. 4. Protodiapha maculifera n.sp., specimen 4/1974, hind' wing.



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318 Psyche [June
ent on the wings, between the veins, and the main veins are covered with a series of setae.
Costa is flattened and in larger wings some- times serrated ; it often carries a series of long setae, which are more pronounced basally. Subcosta consistently forms a conspicuous bend at the very base, probably related to the wing-folding process. Radius is always fused basally with the stem of the media. M, after diverging from R, immediately coalesces or joins either the stem of Cu and CuA, or directly CuA, for a short distance. MA and Rs are con- nected in North American Elmoidae by a cross vein. In Moravian Elmoidae the cross vein is present in only three species. In others, MA almost touches Rs or MA joins Rs at a point, or for a short distance, or both veins are anastomosed for a short distance. Cubitus runs closely to, or touches, R. The stem of Cu is always ribbon-like, deeply concave, broad and flattened. CuA and CUP originate near the point at which M diverges from the common R -I- M stem. Soon after diverging from M, CuA approaches CUP and is connected with it by an oblique cross vein. This feature seems to be a stable character of the family. Anal veins start from the posterior part of the sclerotized region, which is the thickened part of the wing membrane posteriorly from Cu and distally from the basal fold
(fig. 10,s). It undoubtedly served as a reinforcement of the wing base while folding the wings backwards. The basal plate is repre- sented either by the enlarged cubitoanal plate, or by the fusion of the median, cubital, and anal plates. It is partly homologous with the fused subcostoanal plate of the Palaeodictyoptera, the fused basal plate of the Ephemer~~tera, and the radioanal plate of the Odonata. The basal fold runs between Sc and R -I- M, crosses R+ M where it is flattened and weakly sclerotized, follows closely the distal margin of the fused basal plate, and crosses the anal part of the sclerotized region (fig. 1o.B). The axillary sclerite articulates with the proximal margin of the basal plate and with R. Its size and shape are not known.
The family Elmoidae, as long as it was known only by North American species, was well separated from the related Upper Car- boniferous family Diaphanopteridae. The family Diaphanopteridae, represented by Diaphanoptera (France) and Philiasptilon (Soviet Union, Kuznetsk Basin), was characterized by the presence of the anastomosis between MA and Rs. In the Permian material from Moravia, however, this feature is highly variable. In Elmodiapha and Protodiapha (new genera), MA and Rs are connected only by a cross vein, as in the Elmoidae from Kansas and Oklahoma. In the other Moravian genera, MA and Rs are either touching each other



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19741 Kukalova-Peck - Diap hanopterodea
Fig. 5. Protodiapha lineata n.sp., specimen 5/1974, fore wing. Holo- type. Fig. 6. Diapha candida n.sp., specimen 691974, fore wing. Holotype. Fig. 7. Diapha candida n.sp., specimen 7/1974, hind wing. Fig. 8. Per- modiapha carpenter'! nsp., specimen 11/1974, fore wing.



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320 Psyche [June
at a point, or anastomosed as in Diaphanopteridae. Since the pres- ence or absence of the MA-Rs anastomosis is no longer useful as the family character, the status of the Elmoidae as a separate family is questionable. However, since Diaphanoptera, the type genus of the family, is from Upper Carboniferous strata, and since its body structure is entirely unknown, it seems advisable to consider the families distinct at least until more is known about Diaphanoptera. The generic classification used here is based mainly upon the presence or absence of MA-Rs anastomosis, the general shape of the wings, and the character of the Rs area, since these features have been found relatively stable in the related order Palaeodictyoptera ( Kukalovi-Peck, I 97 1 ) .
All specimens available are figured. The lettering of the veins is shown in figures 9 and 10. Genus Elmodiapha, new genus2
Type species: Elmodiapha ovata nsp., Lower Permian of Mora- via.
Fore wing: MA and Rs not anastomosed, connected by a cross vein.
Wings elongated, oval in shape. Sc terminating on R shortly beyond mid-wing. Rs area large, with about 6 branches. R sending off a series of long branches beyond the end of Sc. Hind wing not known.
Elmodiapha differs from Parelmoa in having more elongate wings and richly branched Rs. From Pseudelmoa it differs in the shorter Sc, and in MA approaching more closely to Rs. Elmodiapha ovata n.sp.
Figure I, 2
This species is represented by two detached fore wings with a richly branched Rs.
Fore wing: length 17.5-21 mm, width 5.4-6.6 mm. Costa ser- rated. Rs with 6 long, densely arranged branches. MA simple, MP twice forked. CuA and CUP simple.
IA usually with a series of
branches, anal veins sometimes branched. Holotype: No. 1/1974 ('fore wing fragment, length 18.5 mm, width 6.6 mm, obverse) ; specimen No. 2/1974 (fore wing fragment, length 16 mm, width 5.4 mm, obverse and reverse). Paleontological Institute of Charles University, Prague, Czechoslovakia. 'The latter part of this generic name (and that of the other genera which are proposed in this paper) is derived from the Greek adjective diaphanes (distinct) ; it is treated as a noun and is considered feminine



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19741 Kukalova-Peck - Diap hanopterodea 321 Genus Protodiapha, new genus
Type species: Protodiapha maculifera n.sp., Lower Permian of Moravia.
Fore wing: MA and Rs connected with a very short cross vein. Wings short and broad, broadest at about mid-wing. Posterior margin convexly bent, forming a lobe. Sc terminating on R shortly beyond mid-wing. Rs area small, with about 5 branches. Hind wing similar, but shorter, with the lobe on the posterior margin probably more pronounced.
Protodiapha differs from all the other genera which lack the anastomosis between MA and Rs by the presence of the lobe on the posterior margin.
Protodiapha maculifera nsp.
Figure 3, 4
This species is established upon hind wings having a prominent lobe on the posterior margin and circular, cuticular spots on the wing membrane.
Hind wing: length 14 mm, width 4.5-5 mm. Wing broadest before the level of the end of Sc. The lobe on the posterior margin extending to the end of CUP.
C with a series of long setae. R's
almost straight, with 5 branches.
MA simple, MP with a fork.
CiiA simple, CuP sometimes with short branches. Anal veins with
few branches. Thickened cuticular spots present. Holotype:
No. 3/1974 (hind wing, length 13.5 mm, width 4.5 mm, obverse) ; specimen No. 4/1974 (hind wing fragment, length 8.3 nun, width 5 mm, obverse). Paleontological Institute of Charles University, Prague, Czechoslovakia.
Protodiapha lineata n.sp.
Figure 5 ,
r 7
1 his species is monotypic and based on the proximal half of a fore wing. However, the unique S-shaped cross veins accompanied by dark spots separate the species distinctly from all other Elmoidae. Fore wing: length of the fragment 11.4 mm, width 5.8 mm: Wing broadest beyond the level of the end of Sc. The small lobe- on the posterior margin extending to the end of CUP. Veins darkly
colored. 'CuA with a short branch, CUP simple. Anal veins with few branches. Cross veins long, S-shaped, surrounded by dark spots. Circular cuticular spots absent.




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322 Psyche
Fig. 9. Permodlapha carpenteri n.sp., specimen 8/1974, hind wing Holotype. Fig. 10. Permodlapha carpenteri nsp., specimen 8/1974, hind wing.
Holotype. Ax = axillary sclerite; B = basal fold; FP1 = fused basal plate; S = sclerotized region.




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19 741 Kukalova-Peck - Diaphanopterodea 323 Protodiapha lineata differs from maculifera in having the wings widest beyond mid-wing, a smaller lobe on the posterior margin and long, S-shaped cross veins.
Holotype: No. 5/1974 (fore wing fragment, length 11.4 mm, width 5.8 mm, obverse).
Paleontological Institute of Charles Uni- versity, Prague, Czechoslovakia).
Genus Diapha, new genus
Type species: Diapha candida n.sp., Lower Permian of Moravia. Fore wing: MA anastomosing with Rs for a short distance. Wings slightly broadened beyond mid-wing. Posterior margin con- cave.
Sc terminating on R shortly beyond mid-wing. Rs with 5-6 branches.
Hind wing shorter and probably narrower, with anal veins less branched. Diapha differs from the other genera described herein by the shape of the wings.
Diapha candida n.sp.
Figure 6, 7
A detached fore wing and a hind wing are referred to this species. The hind wing differs slightly in the outline of the posterior margin and in the thinner membrane. It may belong to a different 'species. Fore wing:
length 19 mm, width 5 mm. Wing broader in the distal half.
C basally with a series of more prominent setae. Stem of R distinctly swollen next and distally from the basal fold. Rs concavely bent in the apical part, with 6 branches. MA simple, MP forked. CuA simple, CUP simple or with a branch. IA with several branches, anal veins sometimes branched. Hind wing: length 14 mm, with 4.2 mm. Wing membrane very thin.
Holotype: No. 6/1974 (fore wing, length 18.7 mm, width 5 mm, obverse) ; specimen No. 7/1974 (hind wing, length 14 mm, width 4.2 mm, obverse and reverse). Paleontological Institute of Charles University, Prague, Czechoslovakia.
Genus Permodiapha, new genus
Type species: Permodlapha carpenteri n.sp., Lower Permian of Moravia.
Fore wing:
MA touching Rs or anastomosing with Rs for a short distance. Wing very broad. Posterior margin evenly con- cave. Sc terminating on R shortly beyond mid-wing. Rs with 3-4 branches. Hind wing similar to fore wing, but broader.



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324 Psyche [June
Fig. 11.
Permodiapha lata nsp., specimen 10/1974, fore wing. Fig. 12. Permodiapha &a n.sp., specimen 9/1974. Hind wing, fore wing outlined. Holotype.




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19741 Kukdova-Peck - Diap hanopterodea 325 Permodiapha differs from all other genera with MA-Rs anasto- mosed by its very broad wings.
Permodiapha carpenteri n.~p.~
Figure 8, 9, 10
A detached fore wing and a hind wing are referred to this species. The hind wing No. 8/1974 has been chosen as the holotype because of its well preserved base. The structure of the wing base are discussed above with the family characters. Fore wing: length about 16 mm, width 5.3 mm. Wing large,
with relatively strong membrane, veins and cross veins. Apex prob- ably rounded as in the hind wing. Rs sending off 4 branches, slightly S-shaped distally. MA simple, MP with a long fork. 'CuA simple, CUP simple or forked. Anal veins with several branches. Fore wing about equally long but narrower than the hind wing. Hind wing: length 16.3 mm, width 6.8 mm. Similar to the fore wing but broader, and narrowing more towards the base. Holotype: No. 8/1974 (hind wing, length 16.3 mm, width 6.8 mm, obverse and reverse) ; specimen No. 11/1g74 (fore wing frag- ment, length 11 mm, width 5.3 mm, reverse). Paleontological Institute of Charles University, Prague, Czechoslovakia. Permodiapha lata n.sp.
Figure I I, 12
The species is represented by the holotype, probably a male, con- sisting of a vaguely preserved fore wing, well preserved hind wing and part of the abdomen, and by a detached wing, probably a fore wing.
Fore wing: length about 12.6-15.5 mm, width 4.9-5.2 mm. Wing membrane very delicate, cross veins almost invisible. Apical part of the wing somewhat narrower than in carpenteri. Rs curved posteriorly, sending off 4 branches. MA and CuA simple, MP and CUP with a long fork.
Anal veins with few branches. Fore wing
longer and narrower, with the apical part more pointed than the hind wing. Hind wing: length about 14.5 mm, width 6 mm. An- terior margin strongly curved towards the apex. Permodiapha lata differs from carpenteri in having a very thin membrane and narrower fore wings.
'This species is named for Dr. Frank M. Carpenter (Harvard Univer- sity), who has published extensively on the Diaphanopterodea.



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Fig. 13. Permodiapha specula nsp., specimen 13/1974, fore wing. Holo- type. Fig. 14. Permodiapha sp., specimen 14/1974, hind wing. Fig. 15. Stenodiapha angusta n.sp., specimen 17/1974, fore wing. Holotype. Fig. 16. Stenodiapha angusta n.sp., specimen 18/1974, fore wing.



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19741 Kukalova-Peck - Diaphanopterodea 327 Holotype: No. 9/1974 (fore wing, length about 15.5 mm, width 5.2 mm; hind wing fragment, length 13 mm, width 6 mm; abdomen, I .-10. segment, length 13.5 mm ; hind tibia, length 3 mm ; obverse) ; specimen No. 10/1974 (fore wing fragment, length 12.6 mm, width 4.9 mm, obverse). Paleontological Institute of Charles University, Prague, Czechoslovakia.
Permodiapha specula n.sp.
Figure 13
This species is monotypic and based upon a fore wing. Fore wing: length 13.3 mm, width 4.8 mm. Wing relatively small, with very thin wing membrane. Cross veins almost invisible. Proximal half of the wing much narrower than the distal half. Venation very simple, seldomly branched. MA only touching Rs at a point.
Rs sending off 4 branches, Rs S-shaped distally. MA, MP, CuA and CUP simple. Anal veins mostly simple. Circular cuticular spots present.
Permodiapha specula differs from carpenter! and Zata in having the smallest wings with the most simple venation; in having the wings narrowed in the proximal half; and in the presence of the circular cuticular spots.
Holotype: No. 13/1974 (fore wing, length 13.3 mm, width 4.8 rnm, obverse and reverse). Paleontological Institute of Charles Uni- versity, Czechoslovakia.
Permodiapha sp. specimen No. 14/1g74 (fig. 14) (hind wing fragment, length I I mm, width 4.5 mm, obverse and reverse) might represent a separate species within the genus Permodiapha, but the fragment does not provide enough features for specific diagnosis. Genus Stenodiapha, new genus
Type species: Stenodiapha angusta n.sp., Lower Permian of Moravia.
Fore wing: MA anastomosed with Rs for a short distance. Wing narrow, elongate, with main veins obliquely oriented. Sc terminating on R shortly beyond mid-wing. Rs with 3-4 branches, rs area long and narrow. Hind wing similar to the fore wing, but shorter and broader.
Stenodiapha differs from all other genera with MA-Rs anastomosed by the longest and narrowest wings, with the veins most obliquely oriented.




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Psyche
Fig. 17.
Stenodiapha angusta n.sp., specimen 19/1974, fore wing. Fig. 18. Stenodiapha angusta nsp., specimen 20,4974, hind wing. Fig. 19. Steno- a h a moravica nsp., specimen 21/1974, fore wing, Holotype. Fig. 20. Stenodiapha moravica nsp., specimen 22/1974, fore wing.



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Stenodiapha angusta n.sp.
Figures 15, 16, 17, 18
Three detached fore wings and one hind wing are referred to this species.
Fore wing: length 17.2-20 mm, width 3.8-4.3 mm. Wing broadest at mid-wing, narrowing proximally. Wing membrane very thin. Rs with 4 long branches. MA simple, MP simple; CuA simple, CUP simple or forked.
Anal veins with few branches or simple. Hind wing: length about 16.5 mm, width 4.4 mm. Similar to the fore wing, but shorter.
In hind wing No. 20/1974, the richly branched MP is reminiscent of the Carboniferous genus Diaphanoptera. Holotype: No. 17/1974 (fore wing fragment, length 18.2 mm, width 4.3 mm, obverse) ; specimen No. 18/1974 (fore wing frag- ment, length 15.7 mm, width 3.8 mm, obverse and reverse) ; specimen No. 19/1974 (fore wing fragment, length 14.6 mm, width 3.8 mm, obverse and reverse) ; specimen No. 20/1g74 (hind wing fragment, length 15 mm, width 4.4 mm, obverse and reverse). Paleontological Institute of Charles University, Prague, Czechoslovakia. Stenodiapha moravica nsp.
Figure 19, 20
This species is based on two detached wings, probably fore wings. Fore wing: length about 20.7 mm, width 5.3 mm. Wing broad- ened slightly at mid-wing. C serrated, basally carrying a series of setae. R sending off anteriorly several long twigs beyond th of Sc.
Rs with 3 long branches. MA simple, MP forked. and CUP forked. Anal veins with few branches or simple. Stenodiapha moravica differs from angusta in having the