Jarmila Kukalová.
Revisional Study of the Order Palaeodictyoptera in the Upper Carboniferous Shales of Commentry, France. Part I.
Psyche 76:163-216, 1969.

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REVISIONAL STUDY OF
THE ORDER PALAEODICTYOPTERA IN
THE UPPER CARBONIFEROUS SHALES OF
COMMENTRY, FRANCE. PART I1
BY JARMILA KUKALOVA~
Charles University, Prague
The Upper Carboniferous shales in Commentry, France, are of incomparable significance for the study of insect evolution. The excellent preservation of the fossils and the diversity of insect groups represented make the Commentry fossils basic to any understanding of Palaeozoic insects and early insect evolution. Almost all of the Cornmentry insects are contained in the Institut de Pal6ontoloeie in the Museum National d'Histoire Naturelle, -
Paris, this constituting the largest assemblage in the world of Upper Carboniferous insects. A very few Commentry specimens are in the British Museum (Natural History) in London and in the Man- chester Museum (Stirrup Collection), Manchester, England. The Cornmentry shales are part of a small coal basin, situated on the north side of the large Carboniferous furrow of the Massif Central. The fossiliferous layers are of fresh water origin and are allochthonous, apparently deposited by streams in delta-like, detritic sediments along the shore of a lake.
During his early study on
stratigraphy and sediments ( 1880-1890)) based on surface outcrops of the Cornmentry beds, Fayol assembled the greater part of the collection of fossil insects. This remarkable collection was turned over to Charles Brongniart (grandson of the palaeobotanist, Adolphe Brongniart), who was then an assistant in the Zoological Laboratory of the Museum and who was interested in both geology and en- tomology.
Brongniart's studies were brought out in a single major work, "Recherches pour servir 2 I'histoire des insectes fossiles des temps primaires", published in 1893. This was a pioneer work in the study of fossil insects.
Although his classification of the insects is now seriously out-dated, Brongniart demonstrated in his illustra- tions and his descriptive accounts an exceptional ability for observa- 'Published with the aid of a grant from the Colles Fund of the Museum of Comparative Zoology at Harvard College and a Grant-in-Aid of Re- search from the Society of the
Sigma Xi. This study has also been sup-
ported in part by grants numbered GB2038 and GB7038 (F. M. Carpenter, Principal Investigator) from the National Science Foundation. 'Currently Alexander Agassiz Lecturer in Zoology, Harvard University.



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tion. The collection at the Museum has not been subjected to extensive restudy until recent years. Even Handlirsch, in connection with his monograph of fossil insects in 1906-1908 and his revision of Palaeozoic insects in 1919, made no examination of the Commentry fossils in the Museum. Many additional Commentry insects were found and placed in the Museum subsequent to Brongniart's death in 1899. Most of these were studied and described by Fernand Meunier in a series of superficial and confusing papers ( I 907- I 92 I ) . Professor
Aug. Lameere of Belgium, however, did make a careful study of both the original Brongniart material and the specimens later accumu- lated ; his most significant account ( 1917) dealt with corrections in morphological details and systematics given by Brongniart and Meunier ; his observations were remarkably good, although the com- plete absence of illustrations make the use of his revision somewhat difficult.
In 1935 Professor F. M. Carpenter, realizing both the significance of the Commentry insects and the lack of reliable information about them, decided to make a systematic study of the collection in the Paris Museum in connection with his investigations on Permian insects. He visited the Museum first in 1938, making photographs of all the type specimens, as well as drawings of specimens of some families. Following three other visits to the Museum ( 1961, 1962 and 1966). he published accounts revising the Commentry Protodonata Megase- coptera, Diaphanopterodea, Ephemeroptera, and Caloneurodea. In 1967, realizing the extent of the fossils in the Commentry collection still remaining to be studied, he encouraged me to work on the large order Palaeodictyoptera, a group having more representation in the Commentry collection than in all other collections in the world com- bined. He turned over to me all the photographs and notes which he had previously made, and he obtained financial support from the Scientific Research Society (Sigma Xi) and the National Science Foundation for my visits to the Museum in Paris (1966, 1967) and my work in his laboratory at Harvard University. I am deeply indebted to Professor Carpenter for his assistance, without which the preparation and publication of these studies would not have been possible.
During my stay at the Institut in Paris, I was enabled, through the courtesy of the Director, Professor J. P. Lehman, to restudy all specimens of Palaeodictyoptera in the Museum collection. Dr. J. Sornay of the Institut also kindly gave me great assistance with the collection. Unfortunately, some of the specimens previously present could not be found in 1966 or 1967, the only record of them



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being the photographs made by Professor Carpenter during his stay in Paris before the beginning of World War 11. Although photo- graphic documentation, especially of fossil insects, can be very mis- leading, I decided finally to include figures based on the photographs of these missing specimens, since they provide at least some basis for evaluating the original figures and descriptions given by Brongniart or Meunier.
It is still my hope that the missing types will turn up so that more definitive figures can be made later. For practical reasons, this revisional study will be published in three parts, as follows:
Part I. Spilapteridae (including Lamproptiliidae) , Fouqueidae, Mecynostomatidae, farn. nov.
Part 2. Homoiopteridae, Lycocercidae (including Apopappidae) , Graphiptilidae (including Rhabdoptilidae) , Breyeriidae, Eugereonidae (including Peromapteridae, Dictyoptil- idae) , Ar~haernega~tilidae, Megaptilidae (including Lithoptilidae) .
Part 3. Dictyoneuridae (including Stenodictyidae) . The first two parts will be entirely systematic, but the third will include, in addition to the systematic account of the Dictyoneuridae, a discussion of the morphological features of the Palaeodictyoptera. Since the Commentry insects in the Paris Institut have not been given catalogue numbers, I have followed Professor Carpenter's procedure of referring to Brongniart's specimens by plate and figure numbers as they appeared in his monograph (1893, thesis edition) ; thus, 17-7, refers to the fossil shown in his figure 7, of plate 17. Family Spilapteridae Brongniart
nom. correct. Handlirsch, 1906: 101, pro Spilapterida Brongniart, 1893 : 334.
Spilapteridae, Handlirsch, 1906: 101; Lameere, 1917: 102; Handlirsch, 1919: 20.
Lamproptiliidae, Handlirsch, 1960: 109; Lameere, 1917: 102; Handlirsch, 1919: 21. New synonymy.
Dunbariidae, Handlirsch, 1937 : 81.
Doropteridae, G. Zalessky, 1947 : 64.
Neuburgiidae, Rohdendorf, 1961 : 72.
Type Genus : Spilaptera Brongniart, 1893. This family, the largest in the order Palaeodictyoptera, was estab- lished by Brongniart as a subfamily and subsequently raised to family level by Handlirsch in 1906. Within the order, this family not only has the most extensive fossil record but also the longest range



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I 66 Psyche [June
(Narnurian C to Lower Permian) and wide geographical distribu- tion (North America, Europe and Asia).
Unlike many other Palaeodictyoptera with a tendency to reduce the hind wings, the Spilapteridae and related families always have well-developed hind wings, which are markedly broadened in the basal half. The wing venation is very uniform throughout the family and seems to have been continued into the Lower Permian with rel- atively little change. The detailed branching of the veins, however, is highly variable, even within a single individual. As a result, any classification based upon details of branching of the veins is very questionable.
This study of the Commentry Palaeodictyoptera has brought to light a great deal about the structure of the wings and the body. In the wings transverse structures apparently concerned with strengthening the membrane have turned out to be very characteristic of some genera; since they were diverse, being either cuticular thickenings, oblique cross veins, or oblique, pigmented stripes, they are useful in classification. Many details of body structures not previously noted were observed; these included the fine structures of the antennae, the clvpeus, and the prothoracic lobes. To some extent, their variability within the Palaeodictyoptera is now known. As a result of this more precise and extended knowledge of the structure of the Spilapteridae, I have found it necessary to make some tax- onomic changes. Lamproptilia Brongniart, which is known from fore and hind wings, has a typical spilapterid venation, differing from other genera of the family only in the relatively broad fore wings, the convex curvature of the posterior margin of the hind wing, and the color pattern. None of these features seem to justify more than generic separation. Several generic changes also seem necessary: Co7npsoneura Brongniart, originally referred to the Spilapteridae, is herein placed with some doubt in the family Fouqueidae. Apopappus guernei (Brongniart), also originally placed in the Spilapteridae, is now put in the Lycocercidae. The new genus Tectoptilus is based on Becquerellia grehanti Brongniart. The new species, Homaloneura lehmani, is based upon one of Brongniart's specimens ( I 7- 15 ) , which he erroneously thought was the reverse of his type of Hoinaloneura ornata.
The following are the characteristics which now seem to be valid for the Spilapteridae: wings about equal in length and similar in venation, but the hind wings broader basally; supporting structures often present in basal third of the wings; precostal strip present; anterior margin of wings more or less concave. Sc long, R simple or



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with terminal branches; Rs with 3-10 pectinate branches; both MA and MP with at least two branches, usually more; CuA with several branches, CUP with few branches or even simple; several anal veins. Cross veins mostly simple and not very numerous. The wings are often marked with broad transverse bands or with spots. Body structures: Head small, broad, with large projecting eyes and a large striated clypeus. Beak long; antennae long, with many long segments. Prothoracic lobes with radiating veins and with cross veins, but sometimes heavily sclerotized with the venation reduced or very weak. Metathorax usually somewhat longer than the meso- thorax. Legs short, cursorial. Abdomen relatively slender, the fe- males with ten visible segments and a short ovipositor; cerci in both sexes long and robust. Males of at least some genera with terminal claspers. Posterior-lateral angles of the abdominal tergites small and not projecting.
The family Spilapteridae differs from all other families in the order, excepting the Homoiopteridae (including Rochlingiidae and Thesoneuridae) , Mecynostomatidae, Fouqueidae, and Eubleptidae, in having MA and CuA branched. Of these four families, the Homoiop- teridae show no close relationship to the spilapterids; Mecynosto- matidae, on the contrary, might have been derived from spilapterid ancestors. The Fouqueidae are very close to the Spilapteridae on the basis of their venation, which differs only in the tendency to have fewer branches on MA and more branches on CUP; they have been recognized as a distinct family mainly on the presence of a very dense coarse pattern of cross veins. It might turn out, as explained further below, that the Fouqueidae will be inseparable from the s~ilapterids. The Eubleptidae are apparently closely related to the Spilapteridae (Carpenter, 1965, p. 180) ; the family includes smaller species than those found in the Spilapteridae and is further characterized by having a less developed CuA and the presence of an archedictyon combined with cross veins.
The following Commentry genera are herein included in the fam- ily Spilapteridae : SpiZaptera Brongniart, 'Homaloneura Brongniart, Becquerclia Brongniart, Palaeoptihts Brongniar t, Epitethe Hand- lirsch, Tectoptilus sen. nov., SpiZoptiZus Handlirsch, and Lamprop- nlia Brongniart. Genera which appear to belong in the family, although not occurring in the Cornmentry shales, are the following: Severinopsis Kukalova, 1958 (Namui-ian C, Czechoslovakia) ; Mc- luckiepteron Richardson, I 956 ( Westphalian, Illinois) ; Neuburgia Martynov, 193 I ( Stephanian, Kuznetsk) ; Dmbaria Tillyard, 1924 (Lower Permian, Kansas) ; Oboria Kukalova, 1958 (Lower Per-



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168 Psyche [June
Figure 1. Homaloneura elegans Brongniart; specimen 17-11. Holotype. Figure 2. Homaloneura elegans Brongniart ; specimen 17-12.



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mian Czechoslovakia) ; Doropteron G. Zalessky, 1946 (Lower Per- mian, Ural, USSR) ; Permiakovia Martynov, 1940 (Lower Permian, Ural, USSR) ; Abaptilon G. Zalessky, 1946 (Stephanian, Kuznetsk). The genus Homaloneura occurs in the Westphalian of Illinois as well as in the Cornmentry shales (Carpenter, I 964, p. I I 7ff). Genus Homaloneura Brongniart
Homaloneura Brongniart, 1885: 66; Brongniart, 1893: 316 ; Handlirsch, 1906: 107; Lameere, 1917: 148; Carpenter, 1964: 119. Homaloneurina Handlirsch, 1906: Lameere, 1917 : 148 ; Handlirsch, 1919: 20.
Homaloneurites Handlirsch, 1906: 107; Lameere, 1917: 147; Handlirsch, 1919: 20.
Type species : Hodoneura elegans Brongniart, 1885 (OD). Although Brongniart originally erected HomaZoneura for a single species (elegans), he added five more species in 1893: bonnieri, punctata, joannae, ornata, and buchlandi. In 1906, Handlirsch formed two new genera, their type species being two of those de- scribed by Brongniart : he established Homaloneurina for bonnieri and Homaloneurites for jomnae. Both of these genera were rejected as unnecessary by Lameere (1917)) a view which is undoubtedly correct and which is followed here. An additional species, lehmani, is described below; this is based upon Brongniart's specimen 17-15, which Brongniart erroneously considered to be the reverse half of the type specimen of ornata.
The basic venational pattern within Homaloneura is uniform, and many morphological features seem to have appeared independently among the species. It is usual for the species of Homaloneura to have cuticular thickenings, supporting cross veins, and color bands in the basal third of both pairs of wings. These structures presumably strengthen transversely the thin wing membrane. Among palaeop- terous insects, they are most pronounced in the Odonata, but they occur in other orders as well. In the Palaeodictyoptera, they are most spectacularly represented by the Calvertiellidae (Kukalova, 1964). The strengthening structures in the wings of HomaZoneura were noted originally by Brongniart ( I 893, p. 3 I 8) , but neither Handlirsch nor Lameere made mention of them. In Honzaloneura elegans, bon- nieri parva, and dabaslnskasi, the cuticular thickening is a conspic- uous V-shaped ridge with its apex on AI. In ornata, joannaeo and lehmani, the cuticular ridge is directly on AI, strengthening it for a short distance where it abruptly bends toward the posterior margin. Present in all species is a long, oblique, strong cross vein, running



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1 70 Psyche [June
from M to R, usually at the point of origin of Rs. In elegans and bonnieri, there is additional strengthening by means of a dark colored band running obliquely from AI to R.
The body structures of I!?omaloneura were figured very roughly by Brongniart, and, with the exception of the prothoracic lobes and legs, they were not mentioned by Lameere. Actually, most of these structures are visible only under glycerin or glycerin-alcohol, but with the use of this clearing solution, the specimens of Homaloneura have contributed a great deal to our knowledge of the morphology of the Palaeodictyoptera. Most important is the presence of a large, bulging clypeus, with transverse ridges, best preserved in H. lehmani, which I first noted when I examined this specimen under glycerin in 1966. The structure of the clypeus is surprisingly like that of some Homoptera, such as the Cicadidae, and it undoubtedly indicates that in the Palaeodictyoptera a cibarium was present, much as in the true bugs. This was presumably developed in connection with the sucking beak. In the specimen of lehzani, the sheath of the beak is bent to one side, but the stylets are close together, not separated. Another definite morphological feature of the species of Homa- loneura is the prothoracic lobes.
These were presumably homologous
with wings, but in any event, they show longitudinal veins as well as cross veins, and they are attached to the prothorax along a short, cuticular ridge, corresponding to the articular region of the meso- and metathoracic wings.
The prothoracic lobes are cordate and more or less sclerotized, the veins showing no convexities or concavities. The lobes were somewhat higher on the thorax than the meso- and metathoracic wings, and there was apparently some space between them and the front wings in many species. The size of the lobes varies in the species of the genus. In those species in which the lobes were large, they apparently overlapped to some extent the fore wings. Since there is no indication of articular plates at the base of the lobes, active movement of the lobes in these Palaeodictyoptera is elim- inated. There is a possibility, however, that these lobes might have functioned as vanes in directing the flowing or movement of air in relation to the moving of the front wings. The abdomen in Homaloneura was relatively narrow. Brongniart was of the opinion that only nine abdominal segments existed in the genus Homaloneura ( 1893, p. 3 I 6). This conclusion was appa,rently based on a specimen of ornata, the only Commentry specimen with the abdomen completely preserved, in which segments 1-3 are shorter than the following ones, and in which the posterior margin of the



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second segment is very indistinct. There are actually ten segments visible in this specimen.
In PI. ornata, as well as in many other Palaeodictyoptera, the lateral parts of each tergite are separated by a longitudinal line from the main portion of the sclerite. The structural significance of this is not clear. Carpenter (1969, p. 306) suggests that they might be the actual ventral margins of the tergites, their impressions resulting from the flattening of the abdomen during preservation. On the other hand, they might be homologous with the so-called "lateral lamellae" of some mayfly nymphs, the function of which is apparently uncer- tain.
Description of Genus
Wings equal in length, or the hind pair a little longer. Wing membrane with variable color markings (transverse bands spots, longi- tudinal stripes). Hind wing always broader in the proximal half, but similar in venation to the fore wing.
Cuticular thickenings and r-m
cross vein present. R without terminal branches, Rs pectinate; MA and MP forked several times; CuA with several branches, CUP simple or weakly forked. Cross veins not very numerous. Head about as broad as the prothorax; eyes large, projecting; clypeus large, oval, with median ridges and transverse striations; beak long; antennae long, thin, with long segments. Prothorax nar- rower than and about half as long as the mesothorax; prothoracic lobes cordate, with about ten veins, sometimes branched, and nu- merous cross veins; in some species the lobes are strongly sclerotized and the venation is not visible.
Meso- and metathorax almost equal
in length; legs short and weak. Abdomen relatively narrow, shorter than the wings. Ovipositor short, cerci in the females robust, densely covered by hair; their structure in males is unknown. Species of Homaloneura present in the Commentry shales: PI. elcgans Brongniart, 1885 ; H. bonnieri Brongniart I 893 ; H. punctata Brongniart, 1893; H. bucklandi Brongniart, 1893; and H. lehmani sp. nov. The genus is also represented in the ironstone nodules of Illinois (Westphalian) by PI. dabasinskasi Carpenter, 1964. HomaZuneura elegans Brongniart
Figures I & 2
Homaloneura elegans Brongniart, 1885: 66, pi. 3, fig. 2; Brongniart, 1893: 318, pi. 17, figs. 11-12, pi. 18, fig. 1; HandlirschÌ 1906: 108, pi. 12, fig. 2; Lameere, 1917: 147; Handlirsch, 1919: 20. The photograph of the type specimen was first published by Brong- niart in 1885 and his figure (17-1 I), in 1893. Brongniart referred



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Figure 3. Homaloneura bonnieri Brongniart; specimen 17-17. Holotype. Figure 4. Homaloneura bonnieri Brongniart; specimen 17-18.



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19691 Kukalovh - Palaeodictyoptera I73
to the same species specimen
17-12. Both obverse and reverse of
17-1 I are in the Museum, but only the badly preserved obverse of specimen 17-12 could be found. Fortunately, Dr. Caxpenter put at my disposal a good photograph of the reverse, which he made in 1938, so that a drawing of specimen I 7-12 could be worked out very satisfactorily, also. The following account is based on the type specimen.
Wings slightly unequal, the hind pair being a little longer. Wing membrane very thin. Dark stripes following costa, subcosta, apical margin, and supporting structures. Cuticular thickening V-shaped. Dark band extending from cuticular thickening obliquely to R; a cross vein, rs-ma, running obliquely in the opposite direction. Fore wing: length 33 mm, width 9.3 mm. Anterior and posterior mar- gins almost parallel for about two-thirds of wing length. Apical part narrowed, apex pointed, directed posteriorly. Rs with about 6-7 branches; MA pectinate, with 2-3 branches ; CUP simple. Anal area with about ten branches, sometimes forked. Cross veins few,
almost regular, arranged in two rows parallel with the posterior margin. Hind wing: length 34 mm, width 12 mm. Body structures: Head length 1.5 mm, width 1.4 mm. Eyes of
average size, projecting. Clypeus aln~ost rounded. An tenna.e about 12 mm long, composed of equal, cylindrical segments. Prothorax
half as long as mesothorax, almost square; prothoracic lobe: length 4.7 mm, width 4 mm margin slightly undulated, veins about 11 in number, cross veins not numerous. Mesothorax about 1.4 times longer than metathorax. Fore legs very short, length of femur about 3 mm.
In his description ( I 893)) Brongniart mentions the V-shaped cuticular thickenings and suggests the possibility that they were stridulatory organs, which seems probably incorrect. Homaloneura elegans is related to bonnieri by its thin wing mem- brane, similar color pattern and wing venation. It differs in its smaller size, shape of wings, and the much less dense cross venation of the prothoracic lobes.
Homaloneura bonnieri Brongniart
Figures 3 & 4
Homaloneura bonnieri Brongniart, 1893: 322, 323, text fig. 12, pi. 17, figs. 17-18; Lameere, 1917; 148.
Homaloneurina bonnieri Handlirsch, 1906: 107, pi. 11, fig. 26 ; Handlirsch, 1921: 135, fig. 63.
Brongniart based this species upon two specimens, 17-17 and 17-



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174 Psyche [ J une
18. Since he gave a much more nearly complete figure of specimen 17-17, I am designating it as the lectotype specimen. Handlirsch ( 1906) established the genus ~Homalrincurina for bonnieri without giving any reasons ; Lameere ( I 91 7) was undoubtedly right in synonymizing this genus with Homdoneiira, bonnieri being very close to the type species of the genus.
The type specimen 17-17 shows four rather damaged wings, but I found that the right fore wing could be fully restored by additional preparation and the use of glycerin. Specimen 17-18, which had been badly damaged, I was able to work out much more completely after removal of all the pieces of matrix. Reconstructions of bonnieri have been attempted twice - by Bsongniart ( 1893, p. 323, text-fig. 12) and by Handlirsch ( 1921, p. 135, fig. 63). In both reconstructions, not one restored morphological feature is correctly shown. Specimen 17-18 differs from the type specimen 17-17 in having much larger eyes and in the more convex apical part of the anterior margin of the fore wing, as well as in the more extensively branched MA. Nevertheless, I prefer to consider this to be bonnieri because the large eyes could be due to peculiarities of preservation or to difference in sex, and the other differences mentioned could be due to individual variability. The following account is based upon the type specimen only ( I 7-1 7).
Wings equal in length, the hind pair slightly 'broadened basally. Wing membrane very thin; dark stripes following costa, subcosta, and supporting structures. Cutic~~lar thickening V-shaped. Dark band extending from cuticular thickening obliquely to R. Fore wing: length 43 mm, width 12 mm. Anterior and posterior margins almost parallel. Apex rounded, directed slightly backwards; Rs with about