F. M. Carpenter.
Studies on Carboniferous Insects of Commentry France: Part VI. The Genus Dictyoptilus (Palaeodictyoptera).
Psyche 71(2):104-116, 1964.

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STUDIES ON CARBONIFEROUS INSECTS OF
COMMENTRY, FRANCE: PART VI. THE
GENUS DICT'YOPTILUS (PALAEODICTYOPTERA)* BY F. M. CARPENTER
Harvard University
The genus Dictyoptilus was established by Brongniart in 1893 for a single species, renaulti, based on two small wing fragments. It was placed by Brongniart in the series of Palaeozoic insects which he termed the "Stinodictyopt&ides" and which he assigned to the "Prim- itive Neuroptera". Two very closely related species were subsequently described by Meunier (1908, 1910) in a new genus, Cockerelliella. Although the Meunier specimens are very well preserved, the pub- lished accounts of them by Meunier, Handlii-sch, and Lameere have not included their venational details, which turn out to be very im- portant for the determination of the phylogenetic position of Dic- tyoptih. From a study of these fossils, made at the Laboratoire de PaEontolgie in Paris, in 1938 and 1961, I am convinced that Dic- tyoptil'us is very close to the Permian genus Eugereon and should be included in the family Eugereonidae of the Order Palaeodictyoptera; that the venation of the hind wine of the Eugereonidae has been - -
basically misinterpreted, the pattern being very different from that of the fore wing; and that the wings of Eugereon were actually long and slender, not short and broad, as formerly assumed. The reason" for these conclusions will be given after the descriptions of the Com- mentry fossils belonging to Dictyoptilus. Family Eugereonidae Handlirsch, 1906
Eugereonidae Handlirsch, 1906, Foss. Ins., p. 388 Dictyoptilidae Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23 :I94 Fore wing: slender, costal margin very nearly straight, not arched; Sc long; Rs arising near wing base, with 4 or 5 main branches; stem of M arising independently at base, then aligned with R for a short distance before diverging away; M forking near the level of origin of Rs; MA unbranched; AIP branched; Cu curving towards M R at very base, then parallel to it before dividing; CuA diverging towards M shortly after its origin, unbranched; CUP forked; several *This research has been aided by National Science Foundation Grant No. GB 2038. The previous paper in this series was published in Psyche, 70: 240-256, 1963.




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19 641 Carpenter - Dictyoptilus 105
anal veins arising from one stem. Cross veins numerous, forming a reticulation in some areas of wing.
Hind wing: shape apparently as in fore wing; Rs diverging away from Rl after its origin; base of M apparently independent of R, curved; MA arising at about level of origin of Rs, ending at the position of termination of CuA in fore wing; CUP forking into at least 4 terminal branches; CuA strongly curved. Body structure
(known only in Eugereon) : head small, with slender haustellate beak [For details of body structures see Dohrn, I 867 ; Handlirsch, I 9061.
This family is represented in the Commentry shales by the single genus Dicty optilus.
Genus Dictyoptilus Brongniart
Dictyoptilus Brongniart, 1893, Recherches Hist. Ins. Foss.:390; Handlirsch, 1906, Foss. Ins.:66; Lameere, 1917, Mus. Nat. Hist. Natur., Bull., 23 :I03 Cockcrcllia Meunier, 1908, (non Cockerellia Ashmead, l898), Ann. Soc. Sci. Brux., 32 :I54
Cockcrelliella Meunier, 1909, Ann. Paliont., 4:132 Fore wing: long and slender, the length more than five times the width ; posterior margin with two slight indentations, one near R5 and the other near the end of the posterior branch of MP2. Sc ex-
tending almost to wing apex, RI terminating at very apex; Rs and MA arising at about the same level; Rs with five or more branches; MP dividing into MPI and MP2 directly after its origin; CuA diverging towards M, as characteristic of the family, and either touching or not quite touching &I; cross veins numerous; those in the costal and subcostal areas straight and unbranched; those in other areas straight or reticulate, forming a coarse, irregular network in many parts of the wing.
Hind wing: known only in D. jm-onzapteroides (Meunier) ; shape apparently as in fore wing, with a slightly broader base; space between Rs and MA much broader than that between MA and MP. Body unknown.
Type-species : Dictyoptil,us renaulti Brongniart, I 893 (by mo- notypy) .
The generic characteristics suggested above are tentative, since only one other genus, Eugereon, is known in the family, and since only the basal parts of the wings are known in that genml Dictyoptilus 'The Commentry genus Arc/~aemcgapt'lz~s Meunier ( 1908), which Lameere (1917) considered close to Dictyoptilus, seems to me to require family sep- aration; the wing is relatively broad and lacks the basal divergence of CuA. The family Archaemegaptilidae, established by Handlirsch (1919, p. 13) for this genus, appears to be valid.




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19641 Carpenter - Dictyoptilus 107
seems very similar to Eugereon on the basis of the parts of the wings known in both genera. The basal portions of the fore wings are, in fact, so much alike that generic distinction is not apparent; the hind wings, however, show a few differences, e.g. the area between Rs and MA in Eugereon is fully twice as wide as that of Dictyoptilus. The type-species of Dictyoptilus (renauiti) is known only from two specimens, each consisting of the middle part of a fore wing. However, the venational pattern included is so much like that of the type-species of Cockerelliella (peromapteroides) that the generic syn- onomy given above seems o,bvious.
Dictyoptilus is at present known only from the Commentry shales, in France. Two species (sepultus and per0 mapter oides) have been
described in addition to renaulti. These are quite clearly very close and might well belong to one species. However, since the specific names, which have already become established in the literature, pro- vide convenient means of referring to individual specimens, I am treating them as distinct.
Dictyoptilus renaulti Brongniart
Dictyoptilus renaulti Brongniart, 1893, Recherches. Hist. Ins. Foss.: 391, pi. 22, figs. 13,14; Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23:103. This species was based on the two spec'mens (herein designated 22- I 3 and 22- I 4) figured in Brongniai-t's Recherches, each repre- senting middle portions of a wing. Specimen 22-14 was examined by me at the Museum National, but 22-13 could not be found in the collection. The former is presumably part of a fore wing, on the basis of its ~enation;~ the venation of the hind wing beyond the basal part is unknown in Dictyoptilus.
Brongniart's figure of this fossil is correct, as noted by Lameere 'Brongniartls figure shows a minute fragment of another wing in front of the wing of renaulti, which suggests that the latter was a hind wing. How- ever the venation of specimen 22-14 is not like that of a hind wing, as known in peromapteroides; and the figures in Brongniart's Recherches, although generally accurate so far as the fossils are concerned, are often imaginary with respect to the presence of other fossils on the individual pieces of shale. See Carpenter 1943, p. 529-530.
EXPLANAT'ON OF PLATE 13
Fig. A. Dictyo$tilus sepultus (Mrrnier), original drawing of fore wing based on type in Laboratoire dp Palkontologie, Paris. Fig. B. Dictyoftilus peromapteroides (Meunier) original drawing of hind wing based on type in Laboratoire de Pal6ontologie, Paris.
Lettering: c. costa; Sc. subcosta ; RI. radius; RS, radial sector; MA, anter- ior media; MP, posterior media; CuA, anterior cubitus; CUP, posterior cubitus ; -, concave veins ; "I", convex veins.



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I 08 Psyche [June
( 191 7). The length of the specimen is 32 mm. and its width 19 mm. Comparing this fragment with the corresponding part of the com- plete wing of D. sepultus (Meunier), I estimate that the fragment is about 1/5 the total wing length, indicating that a complete wing of reuauiti would be about 160 mm. long. Dicty optilus sepultus ( Meunier )
Plate I 3, Fig. A; Plate 14
Cockerellia sepulta Meunier, 1910, Ann. Soc. Sci. Brux., 34:195; Meunier, 1910, Mus. Hist. Nat., Bull. 16:235, fig. 3. Cockerelliella sepulta, Meunier, 1912, Ann. Paleont., 7:6; pi. 6, fig. 4, 4a. Dictyoptilus sepultus, Lameere, 1917, Mus. Nat. Hist. Natur., Bull., 23 :160. This species is based on a single, excellent fossil, consisting of a complete fore wing; the veins and cross veins are very clearly pre- served. In one counterpart (the obverse, with Sc concave) the distal third is missing but the rest is exceptionally clear; in the reverse, the basal quarter is missing but the distal portion is very well pre- served. Figure A, plate 13, is a drawing of a complete wing, based on the two counterparts. The total wing length is 106 mm., which is about 50 mm. less than the wing length of renaulti. The wing of
sepultus has a maximum width of about 20 mm. The venation presents no problems in homology, the convexities and concavities being strong- ly indicated. There are two noteworthy aspects of the venation, how- ever.
(I) M arises as an independent vein at the wing base, but shortly diverges anteriorly and continues in contact (but not anas- tomosed) with R for a short distance, forming a double vein; it then separates off as an independent vein.
(2) Cu at its base is directed
anteriorly but shortly runs parallel with R + M, and then divides into CuA ( -I- ) and CUP ( - ) . CuA diverges anteriorly at this point, touching, but not anastomosing, with IVI before diverging away again. These unusual features are duplicated in Eugereon, as noted below.
Lameere (1917, p. 160) has stated that there is a small pi-ecostal space at the base of the wing. A slight thickening of the wing is visible at the base, but I am not convinced that it is actually a pre- costal area. Lameere also states that the subcosta terminates well before the wing apex, as it is shown in Meunier's figure ( I 910) and also in Handlirsch's (1919). Laurentiaux (1957) in an original figure shows Sc extending a little further than indicated in the pre- vious figures. I am convinced from my study of the fossil (as well as of the type of peromapteroides) that Sc extends even further to- wards the apex; at any rate, it is still identifiable as a distinct vein up to that point (See plate 14).




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19641 carpenter - Lktyopti~us 109
hleunier's figure of this fossil ( 1910, p. 236) is unbelievably in- accurate. Handlirsch's illustration ( 19 19, p. 121, although not so crudely done as Meunier's, is erroneous in several major respects) - i.e., short Sc) and the absence of the divergence on CuA. Lauren- tiaux's figure (1953, p. 423) is accurate in all important aspects, but shows a fork on the penultimate branch of Rs, instead of the terminal one.
As noted above, this species is very close to renaulti, but the dif- ference in size is sufficient to retain the species as distinct. Dictyoptilus peromapt,eroides Meunier
Plate 13, Fig- B
Cockerelliu pcromaptcroidcs Meunies) 1908, Ann. Soc. Sci. Bsux., 32 :IS+; Meunier, 1907, Mus. Hist. Natural Bull., 14~36, fig. 2. Cockerelliella pcromupteroides, Meunier, 1909, Ann. Palkont., 4:132) pl. 1, fig. 3.
D;ctyopti/us pcromupteroides, Lameese, 1917, Mus. Nat. Hist. Natur., Bull., 23 :159.
This species is based on a single specimen consisting of a nearly complete fore wing and the basal half of a hind wing; the preserva- tion is satisfactory, although not so good as that of the type of sepuZtlus. The fore wing as preserved is 130 mm. long and has a maximum width of 22 mm. ; the con~plete length of the wing was probably abo~it 14.0 mm. The venation of the fore wing seems to be very close to that of sepultus; in fact, it is difficult to find diffei-ences. The CI-oss veins appear to be a little closer together than those of sepzdtus and the reticulation formed by the ci-oss veins a little finer. The fose wing is about 25 mm. longer than that of sepultus and 30 nm. shorter than that of renaulti.
The type specimen of peromaptemides is especially interesting be- cause of the presence of the hind wing, which is otherwise unknown in Dictyoptdus. 31eunier7s figure of the hind wing (1908, p. 36) is vei-y misleading. Handlissch's figure ( I 91 9, fig. I 3 ) , which was ~nade by a tsacing from Meunier's published photograph of the fossil) is better than Aleunies's but misses many of the important features noted below. Lameere ( 1917) in his brief notes on peronzaj~teroidef makes no comn1ent on the peculiarities of the venation of the hind w~ng.
The hind wing is pseserved only to about the level of the middle of the fore wing; at this point it is clearly bsoken awaj7. There is no indication that the hind wings were substantially shorter than the fore wing) as shown in Handlirsch's figure ( 191 9) ; the distal part of the fragment of the hind wing measures 20 mm. in width) which is 0nl~7 2 mm. less than the fore wing at that position. So far as is



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1 9 64 1 Carpenter - Dictyoptihs 111
known at the present time, therefore, the hind wing was of the same length or nearly the same length as the fore wing. The accompanying drawing (Plate 13, fig. B) of the hind wing of peromapteroides was made from the type specimen in the Museum National in Paris in 1938 and was verified by checking with the specimen in 1961. The
costal margin of the hind wing is not actually visible in the fossil; the basal part is covered up by the hind margin of the fore wing and beyond that point it is broken away; there is) at most, a faint indica- tion of what might be a short piece of the margin just beyond the edge of the fore wing. The first vein which is clearly apparent in the wing is) therefore, the subcosta, which is preserved as a concave vein. Below that) the vein Rl is readily recognizable as a strong convex vein; its basal part is not preserved) being covered by the fore wing. Rs is very closely preserved as a concave vein but unlike Rs in the fore wing, it diverges posteriorly away from Rl and then turns towards Rl; the space between Rl and Rs is actually wider shortly after the origin of Rs than it is further along in the wing. In the part of the
wing preserved, Rs gives rise) in a pectinate manner) to three concave branches, separated by several rows of cellules. Rs in the hind wing) therefore, differs from that in the fore wing by its more basal origin and earlier branching. The rest of the venation of the hind wing is even more different from that of the fore wing. The next vein, which is not obviously convex or concave) arises near the base and is slightly curved; it first gives rise to a strong convex vein, and beyond that it forks to produce two major branches, each in turn forking; this whole system is composed of concave veins. The convex veir* I am identifying as MA, since it follows the distinctly concave Rs; the concave veins below that would appear to be MP. The next and only remaining vein preserved in the wing is a strongly curved, convex vein, apparently CuA; this is not preserved to its termination but the part that is present is almost semicircular. It is difficult to imagine what the distal portion of this hind wing was like; Rs was apparently extensively developed distally, no other main veins remaining. As noted above) there is no evidence that the hind wing was markedly shorter than the fore; the slight indentation of the hind margin cor- responds to the first indentation of the fore wing margin. At any
rate) it is obvious that the fore and hind wings in Dictyoptilus are remarkably different in venation - more so, in fact, than those of any other Palaedictyoptera known. The fore and hind wings of even those genera (as Dunbaria) which show differentiation of wing form, have a similar venation) except for the number and length of anal veins.




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Psyche
The Affinities of DictyoptiZus
As mentioned above) Brongniast? who did not accept Goldenberg's order Palaeodictyoptera) placed Dictyoptilus with several other gen- eral without formal family assignment, in his "N6uropt6res Primaires". Handlirsch ( I 906) included the genus in the family Dict~~oneuridae of the order Palaeodictyoptera. Of course, both Brongniart and Handlirsch knew the genus only by the small frag- ments representing the type-species) renmlti. In I 9 I 7 Lameere? who used a unique system of classification that vistually excluded extinct osders13 erected the family Dictyoptilidae for the genus and five other genera (ArchaenzegaptiZus, Peromaptera, Protagrion, Gilsonia, and Af~ganeura), indicating that he was including the "Pi-otodonates" in the Dictyoptilidae. That familv along with two others (Dictyo- neuridae? Fouqueidae) he included in a major division of the winged insects) the Odonatopt&+es. From this classificationl it is apparent that Lameere regarded DictyoptiZus as more closely related to the obvious odonate types, such as Meganeura, than to the palaeodicty- opterous types? such as Dictyoneura and Stenodictya. In his discussion of the group which he terms the Protodonates ( I 9 I 7, p. I 86) Lameere gives as past of the evidence for this position of DictyoptiZus the presence of a pi-ecostal space at the base of the fore wing and he also points to similarities in venation between Dictyoptilus and such forms as Protagrion, which isl however? now placed in the Pa1aeodictyop- tesa (Carpenter? 1943). As noted above, I am not convinced that there is a true precostal space present in any of the specimens of DictyoptiZus.
Handlirsch in 1919, not accepting Lameere's synonomy of the genus CockereZlielZa with Dictyo~tilus, erected the family C~ckerelliellidae~ which he placed in the Palaeodictyoptera. He assigned Dictyoptihus to the family Dictyoneuridae) as previously. In his posthumously published paper of 1937) Handlirsch reviewed Lameere's classifica- tion, stating his conviction that the Dictyoptilidae of Lameei-e were a mixture of odonate and of true palaedictvoptei-ous types. In this work he recognized the family Dictyoptilidae in the sense of his previous family Cockel-elliellidae? placing it in the Palaeodictyoptei-a. In I 935 Lameei-e still put Dictyoptdus in the Odonoptckes but within the St6nodictoides, thus separating it from the Odonata and Proto- donata (Odonato~ides) .
From the above summary it is apparent that Handlirsch and finally Lameere were convinced that DictyoptiZus was most closely related 3The order Pseudohemiptera (=Protohemiptera) was the only extinct order recognized by Lameere.




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19641 Carpenter - Dicty optilus 113
to the primitive families (e.g.) Dictyoneusidae) of the order Palaeo- dictyoptera. A different view has been advanced by Laui-entiaux (1957)) who associates it with the family Eugereonidae) which has included a single genus from the Permian of Gel-many) and which has become well known because of the presence of a haustellate beak. Eugereon was designated by Handlissch (1906) the type-genus of a new order (Psotohemiptera) but it has been included by most stu- dents of Palaeozoic insects in the Palaeodictyoptera. Lameere ( I 935 ) ) however, separated the group) which he termed the Pseudohemiptera) from the palaeodictyoptesous families (including Dictyoptilidae) by supesordinal lines.
Laurentiaux ( I 953) has elevated the Palaeodic- tyoptera to a superorder and has recognized the Psotohemiptesa as an osdes within that complex. In addition to the Eugeseonidae and Dictyoptilidae) Laurentiaux has included within the Protohemiptesa the Psotagi-ionidae (based on the monotypic genus Protagrion) and the Calvei-tiellidae (based on the monotypic genus CalvertieZZa) . For seasons which will be apparent in the following discussion, I believe Lausentiaux is right in ass'ociating Dictyoptilus with Euqpreon but I do not believe there is evidencle to justify the inclusion of Protagrion and Calvertiella in the same complex.
Text-fig. 1. Eugueon boeckingi Dohrn. A. fore wing; B, hind wing. Original drawings based on photograph of type sent by Dr, Paul Guthorl, Lettering as in Plate 13.
A detailed study of the venation of Dictyo~tilus indicates that this genus is actually closer to Eugereon than has been assumed even by Laurentiaux and) as I have mentioned above, I consider that the two belong to the same family. The basal past of the fore wing) for example, is strikingly similar to that of Eugereon (text-fig. I) ; the stem of $1 arises precisely the same and forms a double vein with the stem of R, although in Eugereon the double vein is somewhat longei- than it is in Dictyoptilus. The stem of Cu arises in the same fashion in both of these genera and CuA diverges anteriorly and touches the



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stem of M. The structure of MA and of MP is essentially the same in the two genera. The anal veins of Dictyoptlius are similar ro those of Eugereon except that they are not quite so strongly curved near the base of the wing. However, it is when we compare the hind wings of these two genera that we find the similarities most striking. Rs, after its origin from Rl, diverges away the same way in the two gen- era and then gives rise to the first of the pectinate branches. The media is curved in both genera and produces the strongly convex MA and after a very short interval MP divides to form the two main concave branches. CuA appears in essentially the same form in both genera.
The dose relationship between Dictyoptilus and Eugereon now seems obvious; the similarities of the fore wings might be due to convergence but the similarities of the remarkably specialized hind wings, even In minor venationd details, make this explanation unten- able. The affinities of these two genera have two interesting implications regarding Eugereon. Since the hind wing of Dtctyoptilus is better and more extensively known than that of Eugereon, our previous interpretations ( Handlirsch, 1906) of the venation of Eugereon now seem to require modification : the vein which has been interpreted as the media in the hind wing is actually the radial sector (Rs) and the former cubitus now turns out to be MA and MP. It might be noted in this connection that Handlirsch's figure of the hind wing of Eugereon ( 1906) shows the very beginning of a branch originating from the vein herein indicated as Rl. Since RI very rarely carries branches in any insect (except distally), Handlirsch apparently identified this vein as the base of the radial sector, which would, of course, be consistent with his interpretation of the next vein as the media. However, no fork or branch of this vein (RI) is shown by Dohrn in his original figure of Eugereon or by any of the other workers who have studied the fossil, and none shows in a photograph of the specimen sent to me by the late Dr. Paul Guthiirl.* Further- more, it is now obvious from the convexities and concavities preserved in the hind wing of DictyoptiIus that the subsequent vein (herein designated as Rs) is a concave vein imd that all of its branches are concave ; if this vein were the media, it should (in the Palaeodictyop- ten) be convex or at any rate have a convex anterior branch (MA). The second implication with respect to Eugereon is the shape of the wings. Although only the basal portions of the wings are preserved 'The figure of Eugcreon included in the Osnovy (B. P. Rohdendorf, 1963, figure 40B) shows the branch on Rl, but that illustration was copied from Handiirsch. 1906.




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19 641 Carpenter - Dictyoptihis 115
in the fossil, Handlirsch has presented a detailed restoration of the entire insect ( I 92 I ) , which shows the complete wing as curiously shaped, short and broad. Haupt (1g4g), basing his conclusions on the same unique specimen, has given another restoration, which shows equally strange but short wings5. Actually, of course, there is no evi- dence whatever for the peculiar shapes of the wings depicted by Handlirsch and Haupt. Indeed, from the similarity of Dictyoptilus to Eugereon, it now becomes virtually certain that the fore wing of Eugereon was long and slender as in Dictyoptilus and that the hind wing was similarly shaped.
Although the Eugereonidae, as conceived here, includes species in which the venational patterns of the fore and hind wings are markedly different, I see no reason fo'r separating the group into a distinct order, as has been done by Laurentiaux. Very little is actually known about most genera of Palaeodictyoptera and as indicated by the history of Dictyoptilus and Eugereon, discussed above, when more informa- tion is obtained, it is usually quite different from what was expected. Attempts to divide the Palae~dict~optera into suborders and super- families, as has been done by Rohdendorf (1961) or into orders, as has been done by Laurentiaux
(1953) seems to me to be useless
nomenclature in the present state of our knowledge of Palaeozoic insects. T'he evidence at hand suggest that the order Palaeodictyop- tera was a very large and diverse group - far more diverse than we have realized - but still monophyletic. It seems highly probable on the basis of the history of other groups of animals that these early winged insects underwent a rapid, radial evolution, but until more structural details are known (e.g., both fore and hind wings, body structure, etc.), I believe we cannot untangle the numerous lines of evolution.
BRONGNIART, c.
1893.
Recherches pour servir A I'histoire des insectes fossiles des temps primaires. Soc. Industr. Minerale, Bull., 7:124-615, pis. 17-53; also published as These Fac. Sci. Paris, 821, pp. 1-494, pis. 1-37. (All
page and plate citations in the present paper refer to the These, since this is the only form of Brongniart's work usually available.)
DOHRN, A.
1866
Eugereon boeckingi. Palaeontographica, 13 :333-340, pi. 41. 1867. Zur Kenntnis der Insekten in der Primarformation von Saar- bru'cken. Palaeont. 14:129-134, pi. 8.
5Both of these restorations are included in Haupt's paper (1949, page 33). The best published photograuh of the actual specimen of Eugereon is in- cluded in Handlirsch, 1906, plate 38.




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Psyche [June
HANDIJRCH, A.
1906. Die Fossilen Insekten, Wien.
1919. Revision der Palaozoischen Insekten, Denkschr. Akad. Wissen- schaften Wien, Math.-Naturwiss. Klass, 96 :1-82. 1921. Palaeonotologie, in Schroder, Handbuch der Entomologie, 3 ~117-306.
HAUPT, H.
1949. Reconstruktionen permokarbonischer Insekten. Beitr. tax. Zool., 1123-43.
LAMEERE, A.
1917. Revision sommaire des insectes fossiles du Stephanien de Com- mentry. Mus. Nat. Hist. Natur., Bull., 23 :141-201 1935. Precis de Zoologic, 4:255-256.
LAURENTIAUX, D.
1953. Classe des insectes, in Piveteau, Trait; de paliontologie, 3 :396-527.
MEUNIER, F.
1908.
Nouveaux Paleodictyopthres du Stephanien de Commentry. Mus. Hist. Natur., Bull. 14 :34-36.
1909. Nouvelles recherches sur les insectes du terrain houiller de Com- mentry. Ann. Palgont., 4:125-152.
1910. Nouveaux Paliodictyoptires du houiller de Commentry. Mus. Hist. Nat., Bull., 16 :233-237.
1910. Une nouvelle esphce de Palkodictyopthre (stknodictyopt6re Brong- niart) du terrain houiller de Commentry. Ann. Soc. Sci. Brux., 34 :195-196.
1912. Nouvelles recherches sur quelques insectes du terrain houiller de Commentry. Deuxihme partie, Ann. Palkont., 7:l-19. ROHDENDORF, B. B.
1962. Osnovy, 9 50-51.




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