F. M. Carpenter.
Studies on North American Carboniferous Insects. 5. Palaeodyctyoptera and Megasecoptera from Illinois and Tennessee, with a Discussion of the Order Sypharopteroidea.
Psyche 74:58-84, 1967.

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STUDIES ON NORTH AMERICAN CARBONIFEROUS
INSECTS. 5. PALAEODICTYOPTERA AND
MEGASECOPTERA FROM ILLINOIS AND TENNESSEE, WITH A DISCUSSION OF
THE ORDER SYPHAROPTEROIDEA.*
BY F. M. CARPENTER
Harvard University
The insects treated in this paper are from the Upper Carboniferous deposits of the "Mazon Creek" region of Illinois and of a new lo- cality in Anderson County, Tennessee. All represent undescribed species, with one exception - the type specimen of Sypharoptera pneuma Handlirsch, for which the author erected a separate order, S ypharopteroidea.
I am indebted to Dr. Sergius Mamay of the U. S. Geological Survey and to Mr. Robert Rapp for the opportunity of studying the Tennessee fossil; and to the authorities of the Peabody Museum at Yale University for the loan of the type of the Sypharoptera. To Mr. Walter Dabasinskas of Cicero, Illinois, I am especially grateful for his kindness in sending me for study more of the remarkable insects which he has collected in the ironstone nodules near Braid- wood, Illinois. The efforts which he and Mrs. Dabasinskas have put into the collecting of fossils in the nodules are a major contri- bution to the study of Carboniferous insects. Family Breyeriidae Handlirsch
This family includes species of large size, having an unbranched MA and CuA, a forked CUP MP with at least three branches and Rs with at least four branches; cross veins are numerous and some- what irregular but do not form a reticulation. The hind wings are broader basally than the fore wings but the venation is essentially the same except for the more numerous anal veins. The family has been found in Upper Carboniferous strata in Eng- land, Belgium, France and Czechosl~vakia.~ The species described below is the first from a North American deposit. The generic *This research has been aided by Grant No. GB 2038 from the National Science Foundation. The previous part in this series was published in Psyche 72: 175-190. 1965.
Pu&e 7458-80 (1967). hup Ytpsychu einclub orgi74/74-058 html



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19671 Carpenter - Carboniferous insects 59 classification of the Breyeriidae has been reviewed by Kukalovi (1959) ; there now seem to be only two valid genera, Breyeria Borre and Strobbsia Handlirsch. The latter is known from the Upper Carboniferous of England but Bre~eria is represented in the other European deposits mentioned under the family and is the genus to which the North American species belongs. Genus BreYeria Borre
Breyeria Borre, C. R. Soc. Ent. Belg., 1875 : 7; synonyms: Borrea Brongniart, 1893 ; Megaptiloides Handlirsch, 1906 ; Pseudoborrea Handlirsch, 1919 ; Breyeriopsis Laurentiaux, 1949.
Breyeria rappi, new species
Plate 8; text-figure I
Fore wing: length 81 mm. ; width 25 mm. Costal margin smooth- ly curved; costal space with a short longitudinal vein basally; costal space not broadened basally; Sc terminating on RI just beyond the first fork of Rs; RI extending to the very apex of the wing; R arising at the level of the fork of M, with six terminal branches; MP with three terminal branches; cross veins mostly straight, a very few branched in the area between Rs and MA; small circular, cuticular thickenings occur between some of the branches of Rs, be- tween Rs and MA, and between MA and MP.
Holotype :
No. I 58550, U.S.N.M., Washington, D.C. ; collected by Mr. Robert Rapp in the "Pewee" Coal Seam at a strip mine near Peach Tree Gap, Anderson Country, Tennessee. The speci- men (Plate 8) consists of a virtually complete wing, lacking only a few millimeters of the apex, and it is perfectly preserved. The species is named for Mr. Rapp. Dr. Mamay has informed me (pers. coin.) that, on the basis of the fossil plants which Mr. Rapp col- lected in the Pewee Coal Seam, he considers the deposit to be of Lower or Middle Allegheny age. This would correspond closely to the Westphalian C Stage of the European Upper Carboniferous. This species differs from others in Breyeria by a series of minor (but presumably specific) characters, such as the general shape of the wing, the number and arrangement of branches of Rs and the pattern of cross veins. The fossil shows no sign of color markings, 'The fossil which Keller (1935) described as Breyeria constantini from the Upper Carboniferous of the Ruhr Valley in Germany is so fragmentary that its assignment to any family is problematical.



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19671 Carpenter - Carboniferous insects 6 I as had been noted by Laurentiaux-Vieira and L,aurentiaux (1964) in Breyeria vrankeni from the Upper Carboniferous of H01land.~ It is most interesting and unexpected to find a representative of the Text-fig. 1. Wing of Breyeria rappi, n. sp. (holotype) family Breyeriidae in the Upper Carboniferous strata of the New World, especially since the fossil shows no differences which separate it generically from the European species. Indeed, the uniformity of the venational pattern of the known Breyeriidae is very striking. The excellence of the preservation of the type of B. rappi reveals some noteworthy features. One of these is the presence of a distinct but short longitudinal vein at the base of the costal space. This is undoubtedly comparable to the several ridges of veins that Kukalovi ( I 960) and Carpenter ( I 964) have noted in other Palaeodictyop- tera. It is by no means certain, however, that this vein is the costa or that a true pre-costal space exists, like that in the Protodonata. Laurentiaux has noted (1964) that he finds no sign of this basal vein in the costal area of the species which he studied. Another interesting feature of the wing of B. rappi is the presence of the small, circular, cuticular thickenings between several of the veins. These have been previously reported in the Breyeriidae by Laurentiaux-Vieira and Laurentiaux ( I 963, I 964). Similar plate-
like thickenings have been noted in some of the Protodonata (Bolton, I 9 I 4, Carpenter I 947 ) and somewhat larger plate-like thickenings The difference in color between the basal and distal parts of the wing of B. rappi, as shown in the photograph, is almost certainly due to peculiari- ties of preservation rather than to pigmentation in the wing of the living insect.
EXPLANATION OF PLATE 8
Breyeria rappi, n. sp. Photograph of holotype. Upper Carboniferous of Tennessee. Length of wing, 81 mm.




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62 Psyche [March
occur in the Diaphanopterodea (Carpenter, 1963) and in some of the orthopteroids. There is no basis, however, for regarding these as nygmata, a term which was used by Navas for certain clear, cuticular spots on the wings of some Trichoptera, Neuroptera and related Endopterygota (see Carpenter, 1963, p. 243, footnote), Family Fouqueidae Handlirsch, I 906
This family has previously been known only from the Upper Car- boniferous shales of Commentry, France, in which it is represented by the genus Fouquea Brongniart and four species. The character- istics which seem to be peculiar to the family are the combination of a branched CuA and very numerous cross veins, which form a true reticulation only along the posterior portion of the wing. The branched CuA, which is not found in the more primitive Palaeodicty- optera, is a characteristic of the Spilapteridae also, but the cross veins in that family are very few in number and are usually aligned to form gradate series.
Mr. Walter Dabasinskas of Cicero, Illinois, has recently sent me a specimen of a palaeodictyopteron which has the same combination of characteristics given above for the Fouqueidae. It cannot, how- ever, be referred to the genus Fouquea and is therefore taken as the basis of the new genus described below.
Genus Neofouquea, new genus
Subcosta closer to R and RI than in Fouquea; CuA deeply forked, i.e., to about the same level as the fork of M into MA and MP; CuA dichotomously branched; CUP deeply forked, without ad- ditional branches. Wings with two slightly curved rows of macula- tions in the basal half; additional maculations probably present in the more distal part of the wing.
Type-species : Neof ouquea suzanneae, n. sp. In contrast to this genus, Fouquea has a pectinately branched CuA and a multiple branched CUP, Cup1 having three or four terminal branches. The cross veins in Neofouquea seem as close together as they are in the other known species of the family Fouqueidae. EXPLANATION OF PLATE 9
Neofouquea suzanneae, n. sp. Photograph of holotype. Upper Carboni- rerous of Illinois.




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64 Psyche
[March
Neofouquea suzanneae, new species
Plate 9 ; text-figure 2
Length of wing as preserved, 30 mm., maximum width, 22 mm.; estimated length of entire wing, based on comparison with species of Fouquea, 65-70 mm. Since only the basal half of the wing, is preserved, the forking of Rs and of MA is unknown, but both of these veins arise much as in the species of Fouquea; MP forks shortly after its origin, MPI being without a branch in the preserved portion of the wing, but MP2 is dichotomously forked at about the level of the outer row of maculations; CuA divides at about the level of the first division of M, much as in Fouquea; CUAI and CuA2 are forked to about the same level; CUP is deeply forked, producing a nearly straight CUPI and a somewhat sigmoidally curved CuPz, as in the other Fouqueidae; the two branches of CUP Text-fig. 2. Wing of Neofouquea suzanneae, n. sp. (holotype). are close together, without secondary branches; the first two anal veins arise from a single stem and the anterior one is forked but not so deeply as in Fouquea. Several anal veins appear to radiate from a basal stem, although their proximal portions are not preserved. The cross veins form an irregular reticulation in the distal portions of the anal veins and along the posterior border of the wing near CUP; in other regions the cross veins are nearly parallel, very close together and only infrequently have connecting branches. An inner



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19671 Carpenter - Carboniferous insects 6 5 row of maculations, consisting of irregularly circular spots, occurs at about the level of the origin of Rs and just beyond the fork of CUP; the more distal row occurs just beyond the level of the fork of MP2 and CuA1 ; there are apparently eight spots in each of the rows.
The type specimen, which is in the collection of Air. Dabasinskas, Cicero, Illinois, consists of basal portions of two wings (see Plate 9). The main veins and the cross veins are well preserved and the macu- lations are clear. The species is named for Suzanne Dabasinskas, in recognition of the persistence and patience which she, as well as her husband, Walter Dabasinskas, have shown in their collecting of the Alazon Creek insects. The specimen of N. suzanneue was col- lected between Braidwood and Coal City Illinois (Will Co.). Since no satisfactory figure of Fouquea Zacroixi Brongniart (the type species of the genus) has previously been published, I include here a drawing of the venational pattern of the type based upon study of the fossil at the Institut de Palbontologie in Paris. A comparison of the venation of Zacroixi with that of suzanneae indi- cates that the type specimen of the latter consists of the basal half of the wing, not quite as far distally as the first branch of the radial sector or of the fork ofMA. The greater width of the wing of the specimen of suzanneae and the curvature of CuA suggest that the specimen is part of a hind wing, not a fore wing; however, since no complete specimen of the family Fouqueidae has been found, we do not have certain knowledge of the difference between the front and the hind wings.
The densely arranged cross veins, which are clearly indicated in the type of suzanneae, are similar to those of the several species of Fouquea in the Cornmentry shales. They are present in the type of Zacroixi although they were not represented in Brongniart's figure of the fossil. They are, however, especially clear over the entire wing of the type of F. superba (Meunier), as shown in the accom- panying photograph (Plate 10) .3
The specimen of Fouquea Zacroixi shows prominent maculations on the wings, arranged in irregular rows, as depicted in Brongniart's figure ( 1893, plate 19, fig. 10). Specimens of the other species of Fouquea [i.e., comstock'i Lameere, needhami Lameere, superba (Meu- 'The genus Archaecompsoneura, which was established for superba (Meunier, 1909), is a synonym of Fouquea, as noted by Lameere (1917)) although this is not apparent from Meunier's grossly inaccurate figures (1910, 1912).




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66 Psyche [March
Text-fig. 3. Wing of Fouquea lacroixl Brongniart ; original drawing based on type specimen in Institut de Palkontologie, Paris. Upper Carboni- ferous of Commentry, France.
nier)] show similar patterns. The maculations on the wings of suzannaea are much smaller and are arranged in definite rows, as shown in the photograph, Plate 9.
The only palaeodictyopteron described from the ironstone nodules of Illinois that resembles N. suzanneae is Diexodes debilis Hand- lirsch ( 191 I ), which was based on a small, poorly preserved wing fragment. However, the cross veins in debilis (the type of which I have examined at the Peabody Museum) are relatively few in number and do not, therefore, form the dense pattern characteristic of the Fouqueidae.
Family Brodiidae Handlirsch, I 906
This family was established by Handlirsch for a single species, Brodia priscotincta Scudder from Upper Carboniferous deposits in England. From the time of its description in 1881, this species has presented numerous problems with respect to both the nature of the structures preserved and their ph~logenetic interpretation. Although numerous specimens (at least 15) have been subsequently found, these have consisted only of isolated wings or fragments of wings; nothing is known, therefore, of the differences between the fore and hind wings or of the body structure. The relationships of Brodia have been uncertain. Handlirsch placed the family at first in the EXPLANATION OF PLATE 10
Fouquea suwba (Meunier).
Photograph of hoiotype, showing pattern of cross veins characteristic of Fouqueidae. Length of wing, 50 mm.



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19671 Carpenter - Carboniferous insects
71
but in all of these the wing base is distinctly petiolate, as shown in Handlirsch's figures (1919). In his diagrammatic representation of the wing, Bolton ( 1921) shows a subpetiolate condition, in which the wing narrows rather more gradually than depicted in Handlirsch's figures. Although there is probably some slight variation in the de- gree of petiolation of the wing, I believe that Handlirsch's figures are more nearly correct in respect to this part of the insect. One of the two specimens in the Scudder collection at the Museum of Compara- tive Zoology shows the petiole distinctly (Plate 11). It is possible, of course, that the fore and hind wings differed in the degree of Text-fig. 4. Comparison of wings of Brodia and Eubrodia. A, Brodia
priscotincta Scudder. Drawing of wing, based on specimens in Scudder Col- lection, M.C.Z. B, Eubrodia dabasinskasi, n. sp. Drawing of wing, based on holotype. The cross-veins and archedictyon are omitted; for these, see plates 12 and 13.
EXPLANATION OF PLATE 11
Brodia priscotincta Scudder.
Upper Carboniferous of England.
Fig.1.
Photograph of specimen no. 5842 in M.C.Z. (Scudder Collection), showing the petiole and the transverse, dark bands on the wing. Actual length of wing as preserved, 55 mm.
Fig. 2.
Photograph of specimen no. 5843 in M.C.Z. (Scudder Collection), showing the dense pattern of fine cross veins; photographed under very oblique light. Actual length of wing, as preserved, 36 mm.



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72 Psyche [March
petiolation but of this we have no information. Bolton figured and described the costal margin at the level of the middle of the petiole as having a distinct hump-like elevation but I have not seen this in any of the specimens which I have examined. There is some uncer- tainty about the shape of the apex of the wing. Curiously enough, this is missing in all specimens of the species which have been indi- vidually described; none of those which were described and figured by Handlirsch and by Bolton show the apex; Bolton includes the apex in his diagrammatic figure of the complete wing but he does not indicate the specimen on which the drawing was based. How- ever, the shape of the wing apex as shown in his "complete wing" is more broadly rounded than that in any specimens which I have seen 01- which have been described by Handlirsch. The wing rep- resented by Laurentiaux (1953) in an original drawing based upon specimens in the Birmingham Museum has a much more bluntly rounded apex; however, his entire figure seems to be somewhat diagrammatic and is probably not intended to represent the precise form of the wing. The absence of the apical part of the wing in so many specimens suggests that the wings of Brodia were unusually fragile and delicate.
(2)
The serration of the wing margins. The serrate margin is one of the more obvious features of Brodia and it has been figured or mentioned in virtually all accounts of this species. The description and especially the figure given by Scudder in 1883 are accurate and detailed. The structure of the costa is very clearly preserved in the specimens in the M.C.Z. (Scudder collection) ; they show that the costa has the serrations along both its anterior and posterior edges, those of the anterior edge being somewhat sharper. Beyond the point where the subcosta joins the costa the serrations along the posterior margin of the costa become more pronounced, as correctly drawn by Scudder (see top fig., Plate 14). The subcosta, which is not actually fused or united with the costa, remains visible as an independent vein. In one of the specimens in the Scudder collection the costal margin is very clearly preserved and shows also in the distal part serrations or short spines on the subcosta in that region, making a total three rows of these short p~ojections.~ It is to be noted in this connection that the radius also bears a few setal bases (see Plate 14). The serrations become very small along the posterior margin distally 'The costal structure of Brodia resembles to a striking degree that of the anisopterous Odonata, in which the costa also has serrations on both the anterior and posterior borders.




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19671 Carpenter - Carboniferous insects 73 but the basal part of the hind margin has a few well-developed ser- rations or heavy stout spines, projecting apically. (3)
The cross veins. These have been the source of considerable confusion. Several distinct cross veins in the pigmented area of the wings have been represented in the drawings of virtually all previous 'a me areas
accounts. They appear to be absent from the light or hj 1' of the wings but I suspect that this is due to the failure of the cross veins to show up in a hyaline area. Bolton was of the opinion, how- ever, that the wing did not, in fact, have color-bands and that the light areas were the result of the wing membrane's being destroyed in those areas. However, the uniformity of the pattern of these dark and light areas in the dozen specimens which have now been found refutes Bolton's theory; furthermore, the presence of the very fine cross veins on the light areas shows that the wing membrane is actually preserved there. The fine indistinct cross veins were noted by Scudder (1883, p. 236) as being "transverse to the nervures they connect, pretty regularly and uniformly distributed over the wing", and Handlirsch ( 1919) represents them in a few of his figures, al- though they were not mentioned by Bolton ( 1921 ). The effect of these numerous, fine transverse veins is to produce a wrinkled ap- pearance on the wing membrane (see figure 2, plate I I ) . They are of special significance in view of the presence of a distinct archedictyon in Eubrodia'; it is my belief that these are vestiges of the archedictyon which are in the process of being lost and replaced by heavier cross veins.
Eubrodia, new genus
Wing: almost certainly petiolate, as in Brodia, but broadest be- yond the middle; pattern of main veins much as in Brodia; no prom- inent cross veins as in Brodia; a fine archedictyon developed over most areas of the wing; in areas just basal and proximal to the mid- wing the archedictyon is absent, the cross veins forming a course reticulation.
Type-species: Eubrodia dabasinskasi, n. sp. Eubrodia dabasinskasi, n. sp.
Plates 12 and 13 ; text-figs. 4B
Wing length as preserved, 48 mm., maximum width, 15 mm.; estimated length of complete wing, 55 mm. The costal margin is slightly concave at about mid-wing, the pterostigmal area being



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19671 Carpenter - Carboniferous insects 75 noticeably convex; apex of wing posterior to the wing axis; Rs with three terminal branches; MP also with three but with one additional terminal twig on MP3 (probably an individual variation); CUP also with a terminal twig. There is a slight, subtle indication of the presence of two light bands across the wing, one just basal of the mid-wing and the other just beyond the mid-wing, these being in