Frank M. Carpenter.
Lower Permian Insects from Oklahoma. Part 2. Orders Ephemeroptera and Palaeodictyoptera.
Psyche 86:261-290, 1979.

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LOWER PERMIAN INSECTS FROM OKLAHOMA
PART 2. ORDERS EPHEMEROPTERA
AND PALAEODICTYOPTERA*
BY FRANK M. CARPENTER
Museum of Comparative Zoology
Harvard University, Cambridge, Mass. 02138 Nature is always on the watch for our follies and trips us up when we strut.-R. W. Emerson
The discovery and exploration of the insect-bearing deposit in the Midco member of the Wellington Formation were made by Dr. Gilbert Raasch and me about forty years ago, just before the begin- ning of the Second World War. Preparation and publication of my first paper on the insects were necessarily deferred until after the war (Carpenter, 1947). By that time I had become convinced of the necessity of my studying in detail as many as possible of the Palaeozoic insects already described from European and North American deposits before continuing with the new material. Having spent several months before the war with the Commentry specimens in the Laboratoire de Palaeontologie in Paris and at least as much time on type specimens in museums in the United States, I had come to realize that many of the published figures and descriptions were unreliable and that most of the fossils had never been properly prepared for study, the body structures usually remaining hidden within the rock matrix. In part because of administrative duties at Harvard University after the war and in part because of the political conditions in Europe during the 1950's, I found it impossible to resume the study of such collections, especially in Paris and Mos- cow, until 1961. Since then I have been able to study the greater part of the more important collections and to publish on some of them, as time and occasion have permitted.
It now seems feasible for me to continue with the series of articles on the insects in the Midco beds. The collection at the Museum of *Partial financial support of this research is gratefully acknowledged to the National Science Foundation, Grant no. DEB-09947, F. M. Carpenter, Principal Investigator. I am also indebted to the authorities of the Peabody Museum, Yale University, for the loan of type specimens.
Manuscript received by the editor October 15, 1979



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262 Psyche [June-September
Comparative Zoology contains about 8,000 specimens from that deposit. Most of them were obtained on the 1940 expedition but others were found from 1948 to 1957. All were collected at the localities listed in Part 1 of this series of papers (Carpenter, 1947). Several years after the publication of that part, Dr. Paul Tasch of the Department of Geology, University of Wichita, Kansas, made several collections of fossils in extensions of the Midco beds or in associated deposits, mainly for the purpose of obtaining Conchos- traca, in which he was especially interested; and with an associate, Dr. J. R. Zimmerman, he published a brief account of some of the insects found there (Tasch & Zimmerman, 1962). I am indebted to Dr. Tasch for placing at my disposal certain of the types in his collection, as well as some unstudied specimens. The previous part of this series of papers dealt with the palaeop- terous orders Megasecoptera, Diaphanopterodea [included as a suborder of Megasecoptera], Protodonata, and Odonata. The pres- ent paper covers the remainder of the palaeopterous orders, the Ephemeroptera and Palaeodictyoptera.
Order Ephemeroptera
Three families of mayflies are known from Permian deposits: Protereismatidae, Misthodotidae (including Eudoteridae) and Pal- ingeni0psidae.l.
The first two of these families are well represented in the Midco beds. Adult mayflies, however, are not nearly as abundant in the Midco deposit as at Elmo, in Kansas. Over a hundred adults have been found in the Elmo beds in collections including about 8,000 specimens; only 26 adults have been found in the Midco beds in an approximately comparable collection. On the other hand, nymphs of mayflies, which are virtually absent at Elmo, are exceedingly abundant in the Midco beds.
Family Protereismatidae Sellards
Protereismephemeridae Sellards, l907:345. Protereismatidae Handlirsch, 19 l9:63
Protereismatidae Tillyard, 1932:237; Carpenter, 1933:489 Kukalovidae Demoulin, 1970:6 (new synonymy) 11 consider this to be a distinct family, not synonymous with Mesephemeridae.



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I 9791 Carpenter - Permian Insects from Oklahoma 263 ADULTS
As now known the potereismatid adults ranged from moderate to large in size. The wings were elongate-oval, without maculations; the hind wings were similar to the fore pair in form and venation, and were only slightly shorter; the costal margin was serrate and prominent setae were present on at least some of the veins; the costal brace was very well developed in both pairs of wings; MA, almost immediately after its origin, coalesced for a short distance with the basal part of RS; RS had three complete triads, and both MP and CUA had a single triad; cross veins were very numerous. The anten- nae, although short, were relatively longer than in existing mayflies; Figure ! (above),
Profereisma direerum, n.sp. Photograph of hol~type (fore wing). MC2 5180a, Permian of Oklahoma. Length of wing, 26 mm. Figure 2 (below).
Prorereisma direcium, n.sp. Drawing of fore wing (holotype). SC, subcosta; R I, radius; RS, radial sector; MA, anterior media; MP. posterior media; CUA, anterior cubitus; CUP, posterior cubitus; cb, costal brace. The con- vexities and concavities of the veins are shown in figure 1.



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264 Psyche [June-September
sclerotized, dentate mandibles were present; the compound eyes were large; all legs were very long and slender, with five tarsal segments; the cerci and the median caudal filament were elongate, and the males possessed prominent claspers. This family, known only from the Permian, was originally de- scribed from the Elmo beds. It is represented in the Midco collection by 18 specimens of adults, as well as by numerous nymphs. All of the Midco specimens belong to the genus Protereisma and most of them to the large and striking species described below. Genus Protereisma Sellards
Protereisma Sellards, l907:347 [For generic synonymy see Tillyard, 1932, and Car- penter, 19331
This genus, the only one at present in the family, is known by five species from Elmo. The insect described by Guthorl(1965) as Prote- reisma rossenrayensis, from an Upper Permian deposit near Rhein- berg, West Germany, is almost certainly a protereismatid, but the published description is not sufficient for generic determination. Two other species, generally referred to as Protereisma uralicum za1essky (1946) (upper part of Lower Permian) and P. apicale (Mar- tynov, 1927) (Upper Permian), both from the Soviet Union, are based on wing fragments that lack parts necessary for family deter- mination. At the present time, therefore, the genus Protereisma is definitely known only from the Lower Permian of Kansas and Oklahoma.
Protereisma directum, n.sp.
Figures 1-4
Fore wing: length 26 mm, width, 6 mm; relatively long and nar- row, the front and hind margins nearly straight; the venation, typi- cal of Protereisma, is shown in figure 2. Holotype: no. 5180ab, collected at locality 15-L, Noble County, Oklahoma, by F. M. Car- penter. This is a complete fore wing, with excellent preservation. The two following specimens are designated as paratypes: no. 5182ab, consisting of the four wings and parts of the body. The fore wing is 31 mm long and 7 mm wide; the hind wing, 28 mm long and 6 mm wide. The body is preserved is dorso-ventral view; the head is 2 mm long and 4 mm wide across the eyes; the prothorax is 1.2 mm long and 4 mm wide; the mesothorax is 3 mm long and 4 mm wide;



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19791 Carpenter - Permian Insects from Oklahoma 265 the metathorax, 2.5 mm long and 4 mm wide. The abdomen (incom- plete) is 25 mm long and 2 mm wide. The other paratype, no. 5 18 1, consists of two fore wings and one hind wing; the fore wings are 28 mm long and 6.5 mm wide; the hind wing, 26 mm long and 6 mm wide.
In addition, there are 11 other specimens apparently belonging to this species, all isolated wings.
The wings of this insect differed from those of other Protereisrna by their large size, slender form, nearly straight anterior and poste- rior margins and the longer costal brace. The species was only slightly larger than P. insigne Tillyard, from Elmo, but the latter had a much broader wing, with a strongly curved posterior margin. P. directurn presumably had a wing spread as great as 70 mm, which is larger than that of most existing mayflies but much smaller than the Jurassic Ephemeropsis tristalis, which had a wing spread of about 90 mm.
Specimen no. MCZ 5182 is of special interest because of the excellent preservation of some parts of the body. The thorax, although somewhat crushed, shows the individual tergites very clearly (figure 4). Previously described specimens of Protereisma from Elmo have shown that the metanotum, although smaller than the mesonotum, was very much larger than it is in existing mayflies; this is shown in the accompanying photograph of directum. The pronotum consisted of a broad plate 1.2 mm long and 4 mm wide, about the same width as the mesonotum.
The serrations along the costal margins of the fore and hind wings of Protereisma were described by Tillyard in 1932. They are clearly visible on the specimens from Midco, especially those of directurn. Tillyard was apparently not aware that the serrations were equally well developed or even more strongly developed on the hind mar- gins of the wings of Protereisma. They are especially well preserved in the neotype of Protereisma latum Sellards, from Elmo (specimen MCZ 34 l9), and I take this opportunity to include two photographs (figure 5) of that specimen here, one showing the serrations along the costal margin and the other, those along the hind margin. The former also shows the setal bases on some of the veins. The serrated margins and setae on the veins are unknown in existing Ephemerop- tera, but they were well developed in the extinct Palaeozoic orders Palaeodictyoptera, Megasecoptera, Diaphanopterodea, and Pro-



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266 Psyche [June-September
Figure 3.
Protereisma direnuin, nsp. Photograph of paratype, MCZ 5182a, Permian of Oktahoma. Length of fore wing, 32 mm. todonata. Serrated anterior margins are present in the existing Odo- nata and occur in isolated families of some other Recent orders, but their functional significance is not under~tood.~ In addition to the specimens of directurn, there are several other isolated wings belonging to Protereisma. One of these (MCZ 5185ab) appears to be a large specimen of P. arcuam Sellards, described from Elmo. Six other specimens are clearly Protereisma but are too incomplete for specific determination. Zimmerman (Tasch and Zimmerman, 1962) has figured a specimen of a mayfly from a deposit a few feet above the Midco insect bed in which the MCZ specimens were collected. I have not seen that specimen, which he identifies as P. hiurn Sellards. It might be that species, but if the costal brace is formed as shown in his figure, the insect could not even be assigned to the Protereismatidae. some existing insects the serrate margins appear to have a function in aggressive behavior. See Owen's account of the butterfly genus Charaxes (1961).



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Carpenter - Permian Insects from Oklahoma 267
Figure 4. Protweisma directurn, nsp. Photopph of head and thoracic region of paratype, MCZ 5182a, showing eyes (E), bead proper (H), pronoturn (Nl), mesono- turn (N2), and metanotum (N3). The left fore wing is preserved with its posterior margin directed anteriorly, as shown in figure 3. The dark circular object to the left of the head is a shell of a conchniitTacan.



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Family Protereismatidae
NYMPHS
In 1968 Dr. Jarmila Kukalova, while making an extended visit to my laboratory at Harvard University, brought from Czechslovakia several fossil mayfly nymphs that she had collected in Permian beds in Moravia. Since only a very few, poorly preserved Palaeozoic mayfly nymphs were known at that time, I suggested that she might also study, along with her specimens from Moravia, some well- preserved specimens that I had collected in the Midco beds in 1940. However, since I had not yet published on or even studied carefully the mayfly adults in that deposit, I requested that the specimens be mentioned by numbers, instead of by new generic or specific names. The reason for that request, of course, was that the systematic posi- tion of the nymphs should be investigated in conjunction with sim- ilar studies of the adults in the same deposit. Accordingly, in Dr. Kukalova's published account (1969) of these nymphs, the fossils were identified as nymphs no. 1, no. 2, etc., of Proterisma sp., the generic assignment being probable but not certain However, my efforts to defer the naming of the Midco nymphs until the adult mayflies had been studied were defeated by Demou- lin with his publication in 1969 of a paper entitled, "Remarques critiques sur des larves 'Ephemeromorpha' du Permien." In this publication Demoulin, without of course seeing any of the speci- mens, formally erected the new genus Kukalova and the new family Kukalovidae to receive the Midco species, which he named ameri- cana (type-species), and one of the Moravian species, moravica. The diagnoses were based on his interpretation of Kukalova's account. He also erected the new genus Jarmila for another of the Moravian nymphs, termed elongata, placing it in the new family Jarmilidae. The two new families were assigned to the extinct order Archodo- nata, and he established a new superorder, Ephemeromorpha, to include the Ephemeroptera and the Archodonata. Had Demoulin communicated his intentions to Dr. Kukalova or to me, we could have corrected his misconceptions of both the nymphs and the Archodonata and thus have prevented the publication of what cer- tainly must be one of the most futile articles in all the literature on fossil insects. That the nymphs from the Midco beds are in fact members of the genus Protereisma will become obvious from the following account. Since the Moravian specimens are not available to me, I am unable to comment on them except by inference.



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19791 Carpenter - Permian Insects from Oklahoma 269 Figure 5. Pratereisma laturn Seliards. Photographs of neotypt, MCZ 341 94 from Permian of Kansas, showing: A, anterior part of hind wing, thearrows pointing to setal bases along the front margin of the wing and on certain veins (X 24); B, posterior part of same wing, the arrows pointing to the serrated hind margin (X38).



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Specimens of the mayfly nymphs are by far the most numerous of all the insects in the Midco deposit. Several hundred were collected on my 1940 trip, when Dr. Raasch and I made the first exploration of the deposit, and as many again were collected on subsequent trips. Double that number were simply discarded in the field. Because of their number and the nature of their preservation, it is virtually certain that these fossils are the cuticular remains shed by the nymphs at molting. More than 90% of the specimens consist of isolated wing-pads from the nymphs and most of the remainder represent a single thoracic segment with two wing-pads attached. Only a very few consist of the entire nymph, with all wing-pads and many body structures, these being the specimens that I turned over to Dr. Kukalova in 1969. Since she has given a detailed account of these specimens, I will include here only the salient features, with special reference to the venation of the wing-pads. The head of the nymphs was slightly narrower than the pro- thorax, and had well developed, dentate mandibles; the antennae were slender; the prothorax about half as long as the mesothorax, and the meso- and metathorax nearly equal; the legs were subequal, with five tarsal segments; abdominal segments subequal, the cerci and caudal filament well-developed; nine pairs of tracheal gills were present on the abdomen, the anterior ones somewhat larger than the others.
The wing-pads were well developed but were attached to the thorax only along the articular area (of the adult wing), and were independent of each other; the pads projected posteriorly at an oblique angle to the longitudinal axis of the body. The venation of the wing-pads was described by Kukalova, but unfortunately her figure (1969, figure 2) and her interpretation of the homologies of the veins were incorrect. The most conspicuous feature of the vena- tional pattern of the nymphal wings is the difference in the apparent degree of development of the convex and concave veins. In the wing of an adult mayfly (figure 8), the convex veins include, in addition to the main veins Rl, MA, and CUA, the intercalary veins of the radial sector and of the posterior media; the concave veins include, in addition to the main veins RS, MP, and CUP, the intercalary veins of the anterior media and the anterior cubitus. In the Midco nymphs (figure 9) all of the convex veins are very strong and distinct but all of the concave veins are weak and indistinct. Comparison of the nymphal wing with the adult wing of Protereisma (figure 8) shows



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19791 Carpenter - Permian Insects from Oklahoma 27 1 Figure 6. Protereisma americanum (Demoulm), nymph. Photograph of holo- type, MCZ 631 1, Permian of Oklahoma. Length of body, 16 mm.



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272 Psyche [.June-September
Figure 7. Proteresstna amencanurn tDemoulin) Photograph of fore and hind wing-pads of holotype. The dark veins are convex, the weak ones (hardly visible) are concave. ~en~th of fore wing-pad, 5.5 mrn. the precise correspondence of the heavy (convex) veins of the nym- phal wing pad with the convex veins of the adult wing, including the intercalary veins of the radial sector and the posterior media. Kuka- lova, in her interpretation of the nymphal wings, apparently as- sumed that all of the heavy veins were the main veins and that all of the weak veins were the intercalary veins. As a result, the true MA was included in her radial sector, the true MP was termed MA, CUA was termed MP, and IA was termed CUP, etc. In figure 9 I include a drawing of the front wing-pad of a nymph (MCZ 8637) with the correct interpretation of the venation, A photograph of the fore and hind wing-pads of the holotype of americana is in figure 7. It is at once obvious from the venation that these nymphs do indeed belong to the genus Protereisma. The presence of the deep fork and triad on CUA eliminates them from the Misthodotidae, for reasons shown below. Demoulin, in removing the nymphs from the Ephemeroptera, was clearly misled by Kukalova's account of their venation but his assignment of them to the order Archodonata was indefensible. The Archodonata had haustellate mouthparts, where- as the nymphs had well developed mandibles. Also, the Archodo- nata lacked the costal brace, as well as the system of triads and intercalary veins, so well developed in the ^In my opinion the Archodonata are members of the order Palacodictyoptera.



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Protereisma americana (Demoulin)
Figures 6, 7, 9 and 10
Kukalova americana Demoulin, 1 97O:6
The holotype specimen is numbered 63 1 1 ab, Museum of Com- parative Zoology; collected by F. M. Carpenter, locality 15-L, Midco insect bed, Noble County, Oklahoma, in 1940 [type desig- nated by Demoulin by reference to plate 29 and figure 1 in Kuka- lova, 19691. This specimen, undoubtedly consisting of the cast cuticle of a nymph, shows the general body structure as well as the four wing-pads. Its dimensions are as follows: fore wing-pad, 5.5 mm long, 1.5 mm wide; hind wing, 4.5 mm long, 1.3 mm wide. The body is 16 mm long, exclusive of the terminal appendages and antennae. A detailed description of this nymph is given in Dr. Kukalova's paper and a photograph of the specimen is included here for convenience of reference (figure 6). This is the best and oldest of the mayfly nymphs that have been found in any Palaeozoic deposit. In addition to the type, there are four other specimens (MCZ 8641- 8644) showing the gills and other characteristic features; all are about 10 mm long, much smaller than the type, and their wing-pads are very small or absent. As noted above, isolated wing-pads are very numerous in the Midco beds. All have the basic pattern of P. americana but of course they may represent more than one species. The smallest (MCZ 8638) of these pads is 2.8 mm long and 1 mm wide (figure 10A); this shows the venational pattern clearly as well as the convexity and concavity of the veins. The largest pad ( MCZ 8636) is 7 mm long and 2.2 mm wide; the cross veins and concave veins are more distinct than in the others (figure 10D). Most of the wing-pads are 5.5 mm long and about 1.7 mm wide (figures 10B, 10C).
There are two aspects of these wing-pads, briefly noted above, that are of unusual interest. One is the distinct fluting of the; pads, even small ones, resulting from the convexity and concavity of the developing veins. The fluting seems to be much more pronounced in Figure 10. Protereisma americanum (Demoulin). Photographs of wing-pads in several stages of development. A, smallest wing-pad found, 2.8 mm long, 1 m m wide, showing definite convexities and concavities; MCZ 8638, Permian of Oklahoma. Lettering as in figure 9. B, wing-pad 5 mm long, seen under oblique lighting; MCZ 8639. C, same specimen as shown in B but with flat lighting, showing the intensity of the convex veins. D, largest wing-pad found, 7 mm long, the concave veins somewhat more distinct; MCZ 8636, Permian of Oklahoma.



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19791 Carpenter - Permian Insects from Oklahoma 275 Figure 10




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the fossil nymphs than in existing ones. It is possible that the greater amount of fluting, which presumably strengthened the wing-pad, may have been correlated with the limited attachment of the pad to the thorax. In this connection it is pertinent to note that wing-pads of the nymphs of the Palaeodictyoptera and Megasecoptera, which also had the limited attachment to the thorax, show a strong fluting. A second feature of interest is the marked difference in the fossil nymphs between the convex and concave veins. The convex veins are preserved as dark brown, thick lines, whereas the concave veins are almost without pigment and appear as fine lines. Even the cross veins (see figure 7) are more obvious than the concave veins. If our inference is correct that these wing-pads represent the cast cuticle of the nymphs, then the dark lines seem to have been pigmented thick- enings on the cuticle that was cast off in molting. I have no explana- tion for the difference in appearance of the convex and concave veins. The pattern of difference is the same in both obverse and reverse halves of the fossils. This eliminates the possible inference that the pattern might have been different on the dorsal as opposed to the ventral surface of the wing-pads. Family Mist hodotidae Tillyard
Misthodotidae Tillyard, 1932: 260
Eudoteridae Demoulin, 1954: 561. New synonymy. The misthodotid adults were of moderate size and generally much smaller than the protereismatids. The wings were broadly oval, usu- ally with maculations, and the hind wings were similar to the fore wings in form and venation, but distinctly broader and with a strongly curved posterior margin. The costal margin was serrate (at least in Misthodotes). The costal brace, although distinct, was weaker than in the Protereismatidae. The venation was basically like that of the protereismatids, except that CUA was unbranched and therefore lacked the triad. Cross veins were somewhat less numerous than in the protereismatids. The body structure is not well known. The antennae were like those of the protereismatids and the mandibles were similarly developed. The legs, however, were apparently much shorter and apparently heteronomous, the fore legs being shorter than the others. The tarsi included four segments (at least in Misthodotes), the 2nd and 3rd being the shor-