F. M. Carpenter.
Studies on Carboniferous Insects from Commentry, France: Part V. The Genus Diaphanoptera and the Order Diaphanopterodea.
Psyche 70(4):240-256, 1963.

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STUDIES ON CARBONIFEROUS INSECTS FROM
COMMENTRY, FRANCE : PART V.
THE GENUS DIAPHAVOPTERA AND THE ORDER
DIAPHANOPTERODEA
BY F. M. CARPENTER
Harvard University
This is the fifth in a series of studies based on the Carboniferous insects from the Commentry Basin, France.l It consists of an analysis of the genus Diaphanoptera Brongniart and a discussion of the Order Diaphanopterodea, which was erected by Handlirsch in I 91 g to receive the genus. In more recent years, there have been described other Car- boniferous and Permian genera which, although previously placed in the Order Megasecoptera, now appear to belong to the Diaphanop- terodea. This group of insects, apparently having a combination of palaeopterous and neopterous characteristics, presents one of the most intriguing and puzzling problems in the geological history of the insects. Our unsatisfactory knowledge of the Commenti-y fossils has added to the difficulties.
Survey of Conzmentry Species
Diaphanoptera was established by Brongniart in 1893 to include two species, D. munieri Brongniart and D. vetusta Brongniart, both from the Commentrv shales. The specimen of one i'munieri) consists of a complete wing, and of the other (vefiusta), of the apical half of a wing. The genus was placed by Brongniart in the group of fossils he termed the "Megasecopterida", including Aspidothorax, Sphecoptera, Psilothorax, etc. In the same publication, Brongniart described a fossil, consisting of a whole but poorly preserved specimen with very long cerci, as Anthracothremma scudderi, placing it in another "family", the "Protephemerides", along with Trifilosoba and Homaloneura. In his 1906 treatise, Handlirsch followed Brongniart's treatment of Diafthanofttera, but he removed scudderi from Afithracothremma, placing it in a new genus, Pseudmthracothremma, which he allocated to an incertae sedis category, the ordinal position being uncertain. 'This research has been aided by a grant (NSF-G14099) from the National Science Foundation and by a previous grant from the Penrose Fund of the American Philosophical Society (1938). I am indebted to the authorities of the Laboratoire de Palt4ontologie of the Museum National d'Histoire Naturelle in Paris for placing at my disposal the unique collections of Commentry insects in the Museum, in 1938. 1961, and 1963 ; and to the authorities of the British Museum (Natural History) for allowing me to examine the Commentry fossils in that institution. The previous paper in this series was published in Psyche, vol. 70. pp. 120-128, 1963.
Fsn-fit 70:HO-256 (1963). http:ffpsyche cnlclub orgfW7O-MO html



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19631 Carpenter - Diaphanoptera 241
Shortly after this, Meunier described ( 1908) as Diaphanoptera su- perba, a specimen which showed all four wings held back over the abdomen ; the venation was very clear but virtually no body parts were preserved. Meunier recognized that the affinites of this fossil were with Brongniart's species of Diaphanoptera,, but he made no comments in his paper on the wings being flexed over the abdomen. Lameere, who examined the Brongniart and Meunier specimens in Paris, noted ( 191 7) that Pseudanthracothremna scudderi is very close to, if not the same species as, Diaphanoptera superba, reaching this conclusion because of the general similarity of size and form of the two fossils, and the nature of the fragmentary venation known in scudderi; and that in both specimens of superba and scudderi the wings rest obliquely along the abdomen (i.e., neopterous-like), not perpendicular to the body (i.e., palaeopterous-like), as in all other Megasecoptera then known. Nevertheless, he continued to place Diaphanoptera in the Megasecoptera. Handlirsch, in his superficial revision of Palaeozoic insects ( 1919), established a new genus, Diaphanopterites, and a new family, Diaphanopteritidae, for Meunier's superba. Unfortunately, he did not see the fossil itself and his interpretation of it was based entirely on Meunier's incorrect drawing and on a small, published photograph. The flexed position of the wings led Handlirsch to re- move these diaphanopterids from the Megasecoptera and to establish a new order, Diaphanopteroidea, for their reception. As a result of my examination of the fossils mentioned above, I propose the following classification of the Diaphanopteridae from the Commentry shales.
Order Diaphanopterodea Handlirsch
Family Diaphanopteridae Handlirsch (synonym : Diaphanop- teritidae Handlirsch)
Genus Diaphanoptera Brongniart (synonyms : Diaphanop- terites Handlirsch ; Pseudanthracothremvia Handlirsch) munieri Brongniart (type-species)
vetusta Brongniart
scudderi ( Brongniart )
superba Meunier
The family Diaphanopteridae is also represented in Upper Cai-boni- ferous strata of the Soviet Union; other families apparently belonging to the Order Diaphanoptei-odea have been found in Upper Carboni- ferous and Permian beds of the Soviet Union and the United States. A discussion of the characteristics and relationships of the Diaphanop- terodea will follow the detailed account of the Commentsv fossils.



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242 Psyche [December
Family Diaphanoptei-idae Handlirsch
Diaphanopteridae Handlirsch, 1906, Foss. Ins. :313 [==: Diaphanopteritidae Handlirsch, 1919 ; Denks., Akad. Wiss. Wien, 96 :65] Fore and hind wings similar; Sc terminating on RI slightly beyond mid-wing; MA diverging away from MP immediately after its origin and just touching or very nearly touching Rs before continuing as an independent, convex vein; CuA coalesced with the base of M. Several large, thickened, circular spots on membrane of both wings. Body Text figure 1. Diaphanoptcra scudden (Brongniart) , after Brongniart, 1893. structure little known; thorax and abdomen combined about as long as wings; abdomen slender; cerci very long, about twice as long as wings.
There seems to me no basis for accepting Handlirsch's family Diaphanopteritidae, which was erected for Diaphanoptcra superha Meunier. The diagnosis given by Handlirsch for the family is very vague and his interpretation of the venation of superba, based entirely on Meunier's published photograph, is inaccurate. In addition to the Genus Diaphanoptera, which is now known only from the Commentry shales, the family Diaphanopteridae is represent- ed in the Upper Carboniferous strata of the Kuznetsk Basin (Asian RSFSR) , Soviet Union, by Philiasptilon maculosum Zalessky ( 193 I ) . Although only the distal half of a wing of this insect is known, its affinites with Diaphanoptera are obvious; two circular spots on the wing membrane correspond approximately in position to spots in Diaphanoptera.




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19631 Carpenter - Diaphanoptera 243
The nature of the circular, cuticular thickenings on the wings of Diaphanoptera is by no means clear. Forbes (1943) identifies them as n~gmata,~ and, incidentally, considers Diaphanoptera to be a true neuropteron, closely related to the living genus Corydalis. However,
cuticular thickenings occur in certain families of Palaeozoic insects which can hardly be regarded as endopterygotes, e.g., the Mischop- teridae of the Megasecoptera, which are clearly Palaeoptera, and the Cacurgidae of the Protorthoptera, to cite only two examples. The
spots in all these Palaeozoic forms are much larger than the nygmata of the endoptei-ygote insects and there is certainly no reason to regard them as homologous structures. Forbes' figure of Diaphanoptera ( 1943) represents the spots as very small, like nygrnata, although they are actually large (see plate 28).
Genus Diaphanoptera Brongniart
Diaphanoptera Brongniart, 1893, Recherches Hist. Ins. Foss. :30S ; Handlirsch, 1.906, Foss. Ins.: 313; Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23 :I+%. Pseudanthracothremma Handlirsch, 1906, Foss. Ins. :324 ; Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23 :148.
Diaphanopterites Handlirsch, 1919, Denkschr. Akad. Wiss. 96 :66. Hind wing very slightly broader distally than the fore wing and apex slightly more rounded. Rs with from j to 7 branches; R4 + 5 parallel with MA; JIA unbranched; MP with from 4 to 5 branches; CuA unbranched ; CUP branched.
Type-species: Diaphanoptera nzunieri Brongniart [Designation by Handlirsch, I 9221.
The generic characteristics given above are somewhat arbitrary, since only one other genus, Philasptilon, is known in the family. The latter, represented by an incomplete wing, differs from Diaphanoptera in having R4 + 5 and IIA converging distally; at the point of its origin R4 + j is much more remote from MA than it is further dis- tally. The rest of the known venation of Philasptilon is very close to that of Diaphanoptera. I am following Lameere in considering Pse.udanthracothrei~z??za a synonym of Diaphanoptera; it has flexed wings and very long cerci, and the few veins that can be seen in the fossil are like those of Diaphanoptera, Diaphanopterites is even more clearly a synonym. The distinguishing characteristics attributed by ^The term nygmata is a modification (Forbes, 1924) of "nigmas" proposed by Navas in 1917 for small cuticular spots which occur on the wings of certain Trichoptera, Neuroptera and related Endopterygota and which grade into similar spots in other insects. Very little is known of their structure and nothing of their function. Martynov (1925) has published the only account of their histology, Martynova (1949) has investigated their presence and distribution in Permian Mecoptera. and Jolivet (1955) has studied the extern- al structure of a variety of types. They have been regarded as probably either glandular or sensory structures.




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19631 Carpenter - Diaphanoptera 245
Handlirsch to the genus da not exist; Sc, for example) was described as extending to the wing apex, but it actually terminates just beyond mid-wing) as in Diaphanoptera.
Diaphanoptera munieri Brongniart
Plate 28, A
Diaphanoptera munieri Brongniart, 1893, Recherches Hist. Ins, Foss. ~309, PI. 17, fig. 10; Handlirsch, 1906, Foss. Ins.:313, PI. 32, fig. 8 ; Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23 :149.
This species was based on a single, well-preserved wing) 38 mm. long and I I mm. wide; on the basis of the shape, I assume it is a hind wing. Its venation is shown in plate 32, fig, A, which is drawn directly from the type specimen in the Laboratoire de PalLontologie. All main veins are clearly preserved and their convexities or concavities distinct. Brongniart's figure of the wing) although very small) correctly repre- sents the venation except in two areas: ( I) The proximal parts of Sc and RI are distinctly bent) as in other primitive Diaphanopterodea; and the stems of R and M are independent) not in contact, as suggested in Brongniart's drawing. (2) MA) distinctly convex) arises from $1 at about the level of origin of Rs and then diverges abruptly towards Rs. These two veins do not, however, quite coalesce) as is shown in Brongniai-t's figure; there is) in fact) a narrow space between them) even at the point of closest association. Handlirsch's figure of C. munieri ( 1~06)~ crudely copied from Brongniart's work)- shows the basal piece of MA as very weak and nearly transverse. This illustra- tion) which is definitely incorrect) has been reproduced in various publi- cations on wing venation and fossil insects (e.g., Cornstock) 1918 ; Rohdendorf, I 962 ) . Forbes) figure ( I 943 ) ) although based on Brong- niart's, is somewhat altered, depicting a broader) mose oval wing) and representing the base of 31A by a broken line. Actually) the basal origin of MA and its divergence to and away from Rs are cIear1y preserved in the fossil.
Like the other diaphanopterids, munieri shows several large spots en the wing. Brongniart indicated six of these in his figure and I find this number in the fossil; but I believe he included one which is actu- &4. Fore wing of D. rn,unieri Brongniart (type). B. Distal part of fore wing of D. vetusta Brongniart (type). C. Fore wing of D. wperba Meunier (type). D. Hind wing of D. superba Brongniart (type). All drawings original, based on specimens in Laboratoire de Paleontologi~, Paris. Sc, subcosta : Rl radius ; R2, R3, R4+ 5, branches of radial sector; MA, anterior media ; MP, .posterior media ; CuA anterior cubitus; CUP, posterior cubitus; +, convex veins; -, concave vems.



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246 Psyche
[December
ally not present and that he omitted one. The one which he omitted is located between CUP and IA; the one which he shows most distally is, I believe) only an irregularity in the rock, not part of the wing; its appearance is very different from that of the others. Diafihanoptera vetusta Brongniart
Plate 28) B
Diaphanoptera vetusta Brongniart, 1893, Recherches Hist. Ins. Fiss.:311; PI. 17, fig. 9; Lameese, 1917, Mus. Nat. Hist, Natur., Bull. 23 1147. This species was based on a well-preserved specimen) consisting of about the distal third of the wing, 20 mm. long and 10 mm. wide; the shape of the apex suggests a fore wing. Brongniart's drawing is essentially correct, except that the first branch of Rs has an additional fork) which he did not show. The species is probably distinct from nzunieri on the basis of the reduced (i.e.) forked) MP and the less extensive Rs. Of particular interest are the wing spots) some of which differ in location fro,m those of nzunieri: there are two between R3 and R4 + 5 and two between MPI and MP2, instead of only one) as in 772 un ieri.
Diaphanoptera scudderi ( Brongniart )
Text figure I
Anthracothremma scudderi Brongniart, 1893, Hist. Ins. Foss. :329 ; PI. 18, fig. 10.
Pseudanthracothre~nma scudderi Handlirsch, 1906, Foss. Ins. :324. Diaphanoptera scudderi Lameere, 1917, Mus. Nat. Hist. Natur., Bull. 23 ~149. This species was based on a poorly preserved fossil) representing a whole insect, the wings (33 mm. long and I I mm. wide) resting
obliquely along the abdomen and the cerci extending fully twice the length of the body; suggestions of the thorax and two legs are present but are too vague to have morphological meaning. Brongniart's figure is essentially correct. The wing venation is so obscure that no satis- factory description or drawing of it can be made ; however) the pattern) so far as it can be seen, is consistent with that of Diaphanoptera, There are faint indications of the wing spots but their disposition is not clear because of the confused venation.
Brongniart) not recognizing the affinities of this fossil with his Dia$hanoptera, placed it in Scudder's genus Anthracothrtmmu, which had been established for an ('orthopteroid" species from North Ameri- ca. Brongniart assigned Anthracothremma to the '(protephemerides'', along with TripZosobu. Handlirsch (1906) correctly removed scudderi from A nthracothremna, erecting a new genus) Pseudanthracothre??~- ma, which he placed ( 1922) in Insecta incertae sedis. In the mean- time, however, Lameere ( 191 7) ) during his examination of the Commentry fossils in Paris) had noted the similarity of the specimen



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19631 Carpenter - Diaphanoptera 247
of scuddcri to the type of lleunier's Diaphanoptera superba and even considered it a possible synonym of sufierba. The significance of the specirnen is that it shows that Diaphanoptera had very long cerci, similar to those subsequently found in other Dia- phanopterodea.
Diaphanoptera superba ATeunier
Plate 28, C, D; Plate 29
Diaphanoptcra superba Meunier, 1908, Ann. SOC. Scient. Brux., 32 :I55 ; 1908, Mus. Nat, Hist. Natur., 'Bull.
14:173; 1909, Ann. Paleont. 4~141, pl. 2, fig. 4; Lameere! 1917, Mus. Nat. Hist. Natur., Bull. 23 ~148. Diaphanopterites superbus Handlirsch, 1919, Denkschr. Akad. Wiss. 96166. The type and 0nl~7 known specimen of this species consists of a whole specimen, shown in ventral view (counterpart not preserved), the wings resting obliquely along the abdomen. The structure of the thorax and abdomen is only vaguely indicated; cerci, as well as other appendages and the head, are not preserved. The wings, however, are very clearly shown and except fox- the parts covered by the abdomen the venation can readily be worked out. Convexities and concavities are distinct. No satisfactory drawing of this fossil has been published. Ifeunier's bears little resemblance to the actual fossil, having sub- petiolate wings and complete absence of Sc in the hind wing. Hand- lirsch's figure, based entirely on Meunier's very small published photograph, is misleading in almost all respects, showing anal lobes on the hind wings. This specimen is the most important of all those known in Diaphanoptera. The drawing in the accompanying figure includes only what can clearly be seen of the venation in the fossil, with some 1-estored
parts indicated by dotted lines. The specimen shows the basal curvature of Sc and RI, as well as the chai-acteristically radiating arrangement of the costal cross veins, In most respects the venation is close to that of nzunieri, although CUP is less branched in the latter. The wing spots are somewhat different; superba lacks the one at the origin of Rs but has an extra one in the area between MP3 and R4+ 5; so far as they are preserved, the spots in the hind wing of superba are placed like those of the fore wing. There are no cerci visible in the specimen of superba. At the end of the abdomen there is a short projection or extension, which might be part of an ovipositor. The Order Diap hanopterodea3
Handlirsch erected this order in 1919 for the fanlily Diaphanop- tesidae. He gave two reasons for the ordinal separation of this family 'Handlirsch's spelling of the ordinal name, Diaphanopteroidea, is unsatis- factory since
the suffix ('oidea" is ordinarily used for subordinal names. I
have accordingly followed Rohdendorf's altered version (1962).



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248 Psyche
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from the Megasecoptera, to which he had previously assigned it: first, the resting position of the wings (along the abdomen, as shown in the type of su~erba) ; and second, the presence of an anal lobe on the hind wing. The latter characteristic does not actually exist in the fossil - Handlirsch simply incorrectly interpreted the photograph of superba published by Meunier. But as to the resting position of the wings, there can be no question.&
The Order Diaphanopterodea has not generally been accepted by students of fossil insects, the Diaphanopteridae being placed in the Order Megasecoptera, as previously. In recent years) however, several families apparently related to the Diaphanopteridae have been de- scribed from Upper Carboniferous strata of the Soviet Union and North America. These fossils, which have in the past been included in the Megasecoptera along with Diaphanoptera, furnish evidence which supports the validity of the Order Diaphanopterodea. In 1961, during my visit to the Institute of Paleozoology at Moscow, I discussed the question of the Diaphanopterodea with the staff of the Institute (Drs. Rohdendorf, Martynova, Sharov, and their associates) and learned that they also were convinced of the validity of the Order Diaphanopterodea. In their subsequent publication, Osnovy ( 1962)- the order is treated as consisting of twelve families. However, since
this work includes no discussion of the reasons for recognizing the order 01- of the general question of its reIationships) I am presenting here my own views on the order and an account of the puzzling mor- phological features of the insects in this group. The following are the families which now appear to belong to the Diaphanopterodea, in addition to Diaphanopteridae : I. Prochoropteridae Handlirsch, I 9 I I (emend. Carpenter, I 940) [Upper Carboniferous, North America].
The genus Prochoropterfi
Handlirsch is based on a single specimen showing the wings held over the abdomen, as in Diaphanoptera, and indicating vague o~ltlines of the abdomen) which bears what appears to be the basal past of a large ovipositor. It was placed by him in the Megasecoptera in 1911 and again in I g I 9, although the Order Diaphan~~terodea was therein erected for Diuphmopteru on the wing position, Haupt (1941) established the Order PaIaeohymenoptera for Prochoroptera but gave no reasons for connecting the genus with the hymenopterous line of insects; like Handlirsch, he did not associate it with the Diaphanop- 4Handlirsch
(1919) treated this difference in wing position as ordinal only and not as indicating a major development in the evolution of insects. Marty- nov (1923) and Crampton (1924) were the first to propose independently the concept of the Palaeoptera 2nd Neoptera (Archipterygota and Neopterygota of Crampton).




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19631 Carpenter - Dia9hanopiera 249
teridae. A second prochopterid, Euchoroptera Carpenter from Kansas (Stanton formation), was based on a single, whole specimen, the wings resting along the abdomen; in addition there is clear preserva- tion of a rostrum (details not discernible), a large ovipositor and a pair of very long cerci, about twice the length of the insect's body. 2. Asthenohymenidae Tillyard, 1924 (emend. Carpenter, 1939) [Lower Permian, Kansas]. This was placed by Tillyard, along with the fa~nily Protohymenidae, in a new order, the Protohymenoptera, but was later transferred to the Megasecoptera. Additional material from Kansas (Carpenter, 193 I, 1939) belonging to both these families showed that although the Protohy~nenidae were palaeopterous (their wings always being preserved in the outspread position), the astheno- hymenids clearly rested with their wings over the abdomen, in an apparently neopterous position. These additional fossils also showed that the asthenomymenids possessed a well developed ovipositor, a pair of very long cerci, and a prominent beak. 3. Martynoviidae Tillyasd, I 932 (emend. Carpenter, I 943) [Low- er Permian, Kansas, Okla.]. Tillyasd placed this family in the sialoid Neuroptera, although he noted some featuses suggesting the Proto- hymenoptera. Additional material, including the hind wings, showed that the martynoviids were close to the Prochoroptei-idae and Astheno- hymenidae (Carpentes, 1947). The body structure and the resting position of the wings are unknown.
4. Elmoidae Tilfyard, 1937 (emend. Carpenter, 1943, 1947) [Lower Permian, Kansas, Okla.]. This was assigned by Tillyard to the Neui-optera, although he recognized a possible relationship to the Protohymenidae and Asthenomymenidae. Additional genera, belong- ing to this family (Carpenter, 1947) from Lower Permian beds of Oklahoma, ~nade the relationship to the Asthenohymenidae and hfarty- noviidae more clear. The body structuse of the Elmoidae is unknown, but a whole specimen shows that the wings were held over the abdo- men at sest.
Using as a basis the features of the five families considered above, we are able to assign the following characteristics to the members of the Order Diaphanoptei-odea: Fore and hind wings hon~onomous or i1early so, the hind wing at most slightly bi-oader than the fore wing, neves with an anal lobe; wings held over or along the sides of the abdomen at rest; head with a prominent beak or rostrum, the detailed structure unknown; cerci very long, fully twice the length of the body. The combination of the rostrunl, flexed wings, and very long cerci is a unique one and certainly justifies ordinal distinction.



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Photograph of ~rpha-napfm mprrhn Mcunier, fvpe specimen (original, X 6), in Laboratoire dc Paleontolngle, Paris. The fine white spots visible on wines and l-iody are mineral in nature and also occur on much of the rock surface.




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19631 Carpenter - Diaphan optera 25 1
Within the order several evolutionary trends are recognizable, of which the following four are the most obvious (see plate 30) : I. The costal area, starting as moderately broad (Diaphanopteridae, El- ~noidae) becomes narrowed distally and eventually very narrow for its entire length (Asthenohymenidae) . This change is correlated with the reduction of the subcosta, which clearly terminates on RI in the Diaphanopteridae but in other families tends to end vaguely in the costal space. The cross veins in the costal space also disappear; in the Diaphanopteridae, Elmoidae, and Martynoviidae, these veins form a definite pattern in the proximal part of the wing, the more basal ones slanting towards the wing base, and the immediate ones following slanting towards the apex. Finally, the membrane between RI and the costal margin tends to thicken, forming a weak pterostigrnal area; this is not visible in the Diaphanopteridae but is in the martynoviids, the pi-ochoi-opterids, and asthenohymenids. 2. The anastomosis of MA and Rs increases greatly. In the Elmoidae the anastomosis between these veins has not even started, but in the Diaphanopteridae it has clearly begun, and the Martynoviidae show a progressive increase which eventually leads to the near loss of the basal part of MA (Phaneroneura) and the ultimate loss in the Asthenohymenidae. 3. The stems of R, M and CuA have become coalesced in the more specialized members of the order. In even the most generalized of these families, the base of CuA has anastomosed with the stem of M, but this compound stem is free from the stem of R (Diaphanopteridae, Elmoidae) ; in the Martynoviidae and Asthenohymenidae, these two stems have fused. In the process of this change, a definite pattern of separation of R, M, and CuA has taken place, this pattern being already discernible in the Diaphanopteridae : R diverges anteriorly from CuA, the angle between them being bisected by the stem of M. This arrangement finally disappears with the loss of the stem of M. It is interesting to note that the basal parts of Sc, R, and M + CuA are arched in all but the most reduced families, this curvature begin- ning in the Diaphanoptei-idae and reaching its maximum development in the Martynoviidae and Asthenohymenidae. .+-
In addition to the families mentioned above, several others, previ- ously assigned to the suborder Paramegasecoptera of the Order Mega- se~optera,~ may also belong to the Diaphanopterodea. These are Parabrodiidae and Raphidiopsidae, from the Upper Carboniferous of the United States; and Kulojidae and Biarmohymenidae from Permian 'The suborder Paramegasecoptera Carpenter was established (1954) for those Megasecoptera which rested with their wings held back over or along the abdomen.




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19631 Carpenter - Diaphanoptera 253
beds in the Soviet Union. At present not enough is known about the structure of the members of these families to permit more definite assignment.
The Permian family Kaltanelmoidae (Soviet Union), placed in the Diaphanopterodea by Rohdendorf (1962), is so little known and its known structure so peculiar that I doubt that it has affinites with the Diaphanopterodea. The Carboniferous family Svpharopteridae (United States), which is also placed in the Diaphanopterodea by Rohdendorf, definitely belongs to another section of the Insecta. Among its other peculiarities is the complete absence of MA, which occurs as a prominent convex vein in the Diaphanopterodea.
Relationships of the Diaphanopterodea
In my account of the Megasecoptera of the Wellington formation in Oklahoma (1947), I discussed in some detail the question of the possible relationships of this order, and in particular of those families in the diaphanopterid-asthenohymenid group. Much of what was presented there now pertains to the relationships of the Diaphanop- terodea. However, two questions now arise in a different form. First, there is the question of the relationship between the Diaphanopterodea and the Megasecoptei-a (s.s.). Tillyard (1936) found it impossible to conceive of the separation of the Asthenohy~nenidae from the Proto- hymenoptera on an ordinal level. However, the evidence now strongly indicates that the sin~ilarities between these two families are entirely a matter of convergence. The coalescence of MA with Rs and of the stem of CuA with M is certainly in this category: a similar coales- cence occurs in several unrelated orders of insects and a great many families within them. What is more important is the distinctive evo- lutionary trend within the Megasecoptera. The tendency for petiola- tion of the wings, for extreme and uniform narrowing of the costal space, the loss of costal cross veins, persistence of setae on the costal margin, the straightness of the stems of Sc, R, and CuA + M - all of these represent significant trends in the Megasecoptera not present in EXPLANATION OF PLATE 30
FOREWINGS OF DIAPHANOPTERODEA
(original drawings)
1. Diaphanoptcru mun'ieri Brongniart, U. Carb., France. 2, Parelmoa revelatu Carpenter, L. Perm., Okla. 3. Martynovia insignis Tillyard, L. Perm., Kans. 4. Eumartynov'a raaschi Carpenter, L. Perm., Okla. 5. Phaneroneura martynovut Carpenter, L. Perm., Okla. 6. Asthenohymen a-picu1i.f Carpenter, L. Perm., Okla.



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254 Psyche [December
the Diaphanopterodea. These, added to the difference in the resting position of the wings, require, in my opinion, ordinal separation. Second, there is the more fundamental question of whether the Diaphanopterodea belong to the Neoptera or Palaeoptera, which for the purpose of this discussion are being regarded as monophyletic groups. Unfortunately, we do not know, and probably never will know, whether or not the mechanism of wing flexing in the Diaphanop- terodea is the same as that in the Neoptera. One possibility, therefore, is that the Diaphanopterodea are true Neoptera. In this case, because of the long cerci and complete venation (including convex I\IA), they should be primitive members of the Neoptera - more primitive, in fact, than any other known order in the series (Protorthoptera, Per- laria, etc.). The difficulty with this theory is the presence of a definite rostrum in the Diaphanopterodea. It is hardly conceivable that the mandibulate trophi of the primitive Neoptera (Perlaria, etc.) were derived from such a specialized type. If the Diaphanopterodea were to be regarded as Neoptera, it would be necessary to assume that they were a specialized derivative of even more generalized Neoptera having mandibulate trophi.
A much more appealing view is that the Diaphanopterodea are direct derivatives of the Palaeodictyoptera and that they developed the wing flexing mechanism independently of the true Neoptera. The venation of the Diaphanopteridae could readily be derived from that of the Palaeodictyoptera and what is more important, the rostrum of the Diaphanopterodea is like that of the Palaeodictyoptera. Actually, the rostrum of Stenodictya ( Laurentiaux, I 952) , which I was able to study in Paris in 1963, is remarkably similar to that of the Astheno- hymenidae.
Until evidence to the contrary is found, therefore, my view of the Diaphanopterodea is that they are phyl~geneticall~ members of the Palaeoptera which have developed a type of wing flexing independently of that of the true Neoptera; and that their closest relatives are the Palaeodictyoptera, from which they were probably directly derived. BRONGNIART, CHARLES
1893 (1894). Recherches pour servir A l'histoire des insectes fossiles des temps primaires. Pp. 1-493.
CARPENTER, F. M.
1931. The Lower Permian insects of Kansas. Part 4. The Order Hemiptera and additions to the Palaeodictyoptera and Protc- hymenoptera. Amer. Journ. Sci. 22 :113-130.



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Carpenter - Diaphanoptera 255
The Lower Permian insects of Kansas. Part 8. Additional Mega- secoptera, Protodonata, Odonata, Homoptera, Psocoptera, Protely- troptera, Plectoptera and Protoperlaria. Proc. Amer. Acad. Arts & Sci. 73 :29-70.
Carboniferous insects from the Stanton Formation, Kansas. Amer. Journ. Sci. 3 8 :636-642.
The Lower Permian insects of Kansas. Part 9. The Orders Neuroptera, Raphidiodea, Caloneurodea and Protorthoptera (Probnisidae), with additional Protodonata and Megasecoptera. Proc. Amer. Acad. Arts & Sci. 75 55-84.
Lower Permian insects from Oklahoma. Part 1. Introduction and the Orders Megasecoptera, Protodonata, and Odonata. Proc. Amer. Acad. Arts & Sci. 76 :25-54.
CRAMPTON, G. C.
1924. The phylogeny and classification of insects. Pomona Journ. Ent. ZOO^. 16:33-34.
FORBES, W. T. M.
1924.
The occurrence of nygmata in the wings of Insecta Holometabola. Ent. News, 3 5 230-232.
1943.
The origin of wings and venational types in insects. Amer. Midi. Natur. 29 :381-405.
HANDLIRSCH, A.
1906. Die fossilen Insekten und die Phylogenie der rezenten Formen. Pp. 1-1430,
1911.
New Paleozoic insects from the vicinity of Mazon Creek. Illinois. Amer. Journ. Sci. 3 1 :297-377.
1919.
Revision der Palaozoischen Insekten. Denkschr. Akad. Wiss. Wien. 9 6 :I-82.
1922. Insecta Palaeozoica (Fossilium Catalogus), (1) I6 :1-230. HAUPT, H,
1940 (1941).
Die altesten geflugelten Insekten und ihre Beziehungen zur Fauna der Jetztzeit. Zeitsch. Naturwissen. Halle. 9450-121. JOLIVET, P.
1955.
Recherches sur les organes facettiques des ailes des insectes. Inst. Roy. Sci. Nat. Belg. Bull. 3 1 :1-23.
LAMEERE, A.
1917. Revision sommaire des insectes fossiles du Stephanien de Com- mentry. Mus. Nat. d'Hist. Natur. 23 :141-201. LAURENTIAUX, D.
1952, Dkouverte d'un rostre chez Stenodictya lobata Brgt. (Palkodicty- optere stknodictyide) et Ie problkme des Protohemipteres. Bull. Soc. geol. France (6) 2 233-247.
MARTYNOV, A. V.
1923.
On two basic types of insect wings and their significance for the general classification of insects. Trudy 1st All Russ. Congress 2001. Anat. Histol. (Petrograd), 1922 38-89. [Russianl. 1925.
On the facetic organs on the wings of insects. Trav. Soc. Natur. Leningrad 44 :1-23.
MARTYNOVA, 0.
1949. Facetic organs on wings of Mecoptera. Bull. Mosc. Obsh. Isp. Priroda, Geol. sec. 24:93-95. [Russian]. MEUNIER, F.
1908.
Deux nouveaux Migas6copt6res st un nouveau Pal6odictyopte1-e du Stephanien de Commentry. Ann. Soc. Sci. Brux. 322-3. NAVAS, L.
1917. Algunos organos delas alas de 10s insectos (4). Asoc. Espafiola para el Progreso de las Ciencias, Pp. 57-62.



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256 Psyche [December
ROHDENDORF, B.B., et al.
1962. Osnovy Paleontologii. Akad. Nauk USSR, pp. 1-560. [Russian]. TILLYARD, R. J.
1924, Kansas Permian insects. Part 3. The New Order Protohymenop- tera. Amer. Journ. Sci. 8 :Ill-122.
1926. Kansas Permian insects. Part 16. The Order Plectoptera (contd) : the family Doteridae, with a note on the affinities of the Order Protohymenoptera. Amer. Journ. Sci. 3 2 :435-453. 1937. Kansas Permian insects. Part 17. The Orders Megasecoptera and additions to the Palaeodictyoptera, Odonata, Protoperlaria, Cope- ognatha and Neuroptera. Amer. Journ. Sci. 33 : 81-110.



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