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The Lower Permian Insects of Kansas. Part 11. The Orders Protorthoptera and Orthoptera.
Psyche 73(1):46-88, 1966.
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THE LOWER PERMIAN INSECTS OF KANSAS. PART 11. THE ORDERS PROTORTHOPTERA
AND ORTHOPTERA1
BY F. M. CARPENTER
Harvard University
The two preceding papers in this series dealt with representatives of seven closely related families of the Order Protorthoptera occurring in the Elmo limestone.
The present paper treats additional families of more diverse relationships within that order and also covers several families of the Order Orthoptera.
The problems involved in the systematics of the Palaeozoic orthop- teroids are intrinsically very great, mainly as a result of our frag- mentary knowledge of most species but also as a result of the variability of the venation within species. It was my belief more than twenty years ago (1943, pp. 76-77) that the classification of the Palaeozoic orthopteroids as suggested by Handlirsch first and by Martynov later was not realistic in the light of our knowledge at that time.
Since then many additional orthopteroids have been de- scribed, mostly from the Lower and Upper Permian strata of the Soviet Union. These new fossils have added greatly to our knowl- - -
edge of the early history of the orthopteroid complex. Through the courtesy of Dr. B. B. Rohdendorf, Arthropod Section, Palaeontolog- ical Institute. Academy of Sciences, in Moscow, I had the opportunity in 1961 of studying both the undescribed and described material in the collection of the Institute; and of discussing with Dr. Sharov, Dr. Martynova, Dr. Bekker-Migdisova and other staff members of the Institute various oroblems of insect evolution. I would be remiss if I did not acknowledge at this time my gratitude to the entire staff of the Institute for their kindness and help during and subsequent to my stay.
During the past decade I have been able to study additional orthop- teroids collected at the Elmo locality and especially in the Midco beds in Oklahoma. Two additional trips to, the Institute de Palionto- logic in Paris have enabled me to make further examination of the Commentry fossils, which I still consider (in spite of the remarkable fossil insects from Tchekarda in the Soviet Union) the foundation on which our understanding of Palaeozoic insects rests. 'This research has been supported in part by a National Science Founda- tion Grant, No. GB 2038.
Part 10 of this series was published in the Proc. Amer. Acad. Arts Sci., 78 ~185-219, 1950.
Pnche 73:46-88 (1066). hup //psyche enlclub orgi73173-IM6 html
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19661 Carpenter - Protorthoptera and Orthopiera 47 In the present state of our knowledge, the classification of the Palaeozoic orthopteroids is necessarily based on the venation of the fore wings, the hind wings and body structures being very little known at best and entirely unknown in by far the majority of species. Since considerable difference of interpretation exists in even the re- cent literature on the orthopteroid venation, I consider it necessary to present here my own views on the homologies of wing veins in these particular insects and indeed in insects in general. I find that few students of insects have any understanding of the problems of vein homology or of the current status of the subject. The following account is intended to present the background and the nature of my own views used throughout this paper and the subsequent parts in the series.
Although some preliminary attempt was made by Hagen (1870) to hokologize the wing veins of insects, Red tenbacher ( I 886) was the first to make a significant contribution to the subject. He pro- posed the recognition of six main veins, which he termed the costa, subcosta, radius, media, cubitus and anal. In reaching his conclusions, he considered the general correlation of the positions of the veins, as well as a primitive alternation of topography, i.e., convexity and concavity. The Redtenbacher System of nomenclature was followed by Comstock and is actually the one which has been in general use, although it is commonly referred to as the Comstock-Needham Sys- Comstock's first publications on wing veins appeared in 1892. In 1895, J. G. Needham, then a graduate student under Comstock, began a new approach to the study of wing vein homology and the ontogenetic development of wings and their veins. Results of these studies were first published in a series of articles under joint author- ship of Comstock and Needham in 1898 and 1899. An extensive series of papers, mainly by Needham, appeared in subsequent years and in 1918 Comstock brought together in book form a compilation of what had been done in his and Needham's laboratories. They con- dueled that the various patterns of wing venation in insects had been derived from a common ancestral type and that the veins of different orders could be homologized. The Redtenbacher System of nomen- clature was used by them, although no significance was attached to the convexity or concavity of the veins. As noted above, the innovation brought into their venational studies was the ontogenetic method. Noting that in such primitive insects Tomstock himself pointed out (1918, p. 11) that this nomenclature should be recognized as the Redtenbacher System, not the Cornstock-Needham System.
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48 Psyche [March
as the Plecoptera only tracheae could be seen in the developing wings, and that the pattern of venation of the adult wing agreed closely with the pattern of tracheation in the wing pad, they concluded that tracheae determined the positions of the veins (Comstock, 1918, p. 12). They also concluded that the ontogenetic history of the tracheal pattern recapitulated the phylogenetic history of the vena- tion in the group of insects concerned. Applying these principles to the Odonata and Ephemeroptera, for example, they reached the un- expected conclusion that a branch of the radius vein had crossed over a branch of the media in the course of the evolution of these groups; the trachea appeared to cross over in the wing pad and this, in their view, meant that the vein had done likewise in previous geologic time. Objections to the tracheation theory of vein deter- mination and especially to the recapitulative conclusions were raised by several students of fossil insects and insect evolution (e.g., Till- yard, Martynov, Carpenter, Fraser ) in the period from 1923-1935. In 1935, Needhain reiterated his stand on the ontogenetic-phylogenetic relationship of tracheae and veins ; and in I 95 I, he published a more detailed discussion in defense of this thesis, especially as it related to the Odonata; although a substantial part of his paper was an at- tempt to ridicule in a personal manner all individuals who had dis- agreed with him.;
As Needham himself indicated ( 1935, p. 129) there had not been undertaken up to that time a thoroughgoing investigation of the de- velopment of nymphal wings of any species, at least with respect to the development of tracheae and veins. Shortly after, however, such an investigation was made by Holdsworth ( I 940, 1941 ) , this con- sisting of a histological study of the development of wing pads, tracheae and veins, starting with the earliest beginnings of the wing buds. The plecopteran, Pteronarcys, was chosen because Comstock and Needham considered the stone-flies as demonstrating most clearly the tracheal determination of veins. Holdsworth's results were strikingly clear: the tracheae did not enter the main area of the wing pads until the blood spaces or lacunae between the blocks of epidermal cells had already established the positions of the veins. The tracheae, as they grew longer, simply entered the lacunae which had already been blocked out, following the lines of least resistance. Variation in the tracheal branching was obvious and usually several Qne can only regret that this final paper on this subject by Needham was so vindictive. It contributed nothing to science and detracted from Needham's image as a scientist. It also earned a black mark for the Amer- ican Entomological Society for publishing it.
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19661 Carpenter - Protorthoptera and Orthopiera 49 lacunae received no tracheae. Eventually, the epidermal cells lining [he lacunae, including those without tracheae, secreted the cuticular materials which finally formed the veins. The obvious conclusion from this investigation was that the tracheae did not determine the positions of the veins. What Comstock and Needham had observed was the entrance of the tracheae into the wing pad, followed by vein formation, which ultimately closely resembled the tracheal pattern. What they did not see was that the blood lacunae, along which the veins would form, were already blocked out, before the development and extension of the tracheae.
Holds-worth's conclusions have been corroborated by the investiga- tions of Henke ( 1953) and of Leston ( I 962) on the inter-relation- ships of veins and tracheae, demonstrating that the lacunae in wing pads aze the precursors of veins, the tracheae merely occupying the available lacunae. Smart ( 1961) has shown that the cutting of the main tracheae in the wing pad of Periplaneta resulted in degeneration of tracheal branches and in retracheation but with an abnormal pat- tern, which, however, had no effect on the normal venational pattern. His conclusion was that the pattern of tracheation of the nymphal or the pupal wing could not be taken as fundamental in determining the homologies of the veins.* As the situation now stands, the Comstock- Needham method ot determining homologies of veins, which domi- nated investigations of wings for the first half of the present century, must be regarded as a side issue which actually led nowhere. How- ever, it must also be emphasized that many of the conclusions reached by Comstock and Needham, not involving their ontogenetic method, are perfectly valid.
Another approach to the problem of homologies was introduced by Lameere in 1923, as 2. result of his extended and important studies on the Carboniferous insects of Cornmentry, France. Impressed by the regularity of the convexities and concavities, he concluded that there were originally two media veins and two cubitus veins, one of each being convex ( + ) and the other concave ( - ) ; these he termed the media anterior (MA), media posterior (MP), cubitus anterior (CUA) and cubitus posterior (CUP). He believed that some insects had both convex and concave elements, while others had various combinations of one or the other. Support? for his con- " have given this detailed summary of the Comstock-Needham method of determining wing homologies because their conclusions, based on this tech- nique, have become firmly implanted in American entomological literature and in current texts. See, for example, the 1963 edition of Borror and DeLong's "An Introduction to the Study of Insects."
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50 Psyche [March
clusions has come from the study of Palaeozoic insects and more primitive groups of living insects with the result that the Lameere view has been generally accepted as a working hypothesis by stu- dents of fossil insects and insect evolution. In this connection, one should recall that Redtenbacher in his original account of vein homologies used the alternation of convexities as part of the evidence for his system of homologies. Unfortunately, the convexity or con- cavity of several veins has been lost in most orders of insects. The subcosta ( - ) , radius ( + ) , radial sector ( - ) , anterior cubitus ( 4- ) ,md posterior cubitus ( - ) tend to retain their topography in mem- branous wings, although virtually all veins in thick tegmina or elytra appear to have lost their topographic positions. The anterior media ( + )
and posterior media ( -) have generally come to lie flat in the wing membrane, except in the palaeopterous orders, where they are distinctly different. As a working hypothesis, I am assuming the presence of both of these veins in the early neopterous stock; there is some evidence from the pattern of these two veins in closely related taxa that they have been retained even in the endopterygote line (see, for example, Carpenter, 1940, Adams, 1958). Histological investigations on the development of convex and concave veins are still needed. Holdsworth included some histological observations in his work previously cited, but his studies were limited to one species. Mayfly wings, treated with caustic potash, separate into their orig- inal membranes, all the convex veins being on the dorsal membrane and all the concave veins on the ventral membrane (Speith, 1932 ; Holdsworth, 1941). Holdsworth noted that, although there was not this sharp difference in Pteronarcys, most of the cuticular material of the convex veins appeared to be formed in the dorsal epidermal layer and most of that of the concave veins in the lower epidermal layer.
It is not improbable that this is generally the case. As noted above, veins have tended to lose the topographic characteristics in tegmina or elytra; and it is possible that a previously concave vein might eventually acquire a convex position secondarily if the tegmen became membranous. However, I regarded the latter occurrence as probably a rare event and consider convexities or concavities of veins as due to the original condition, unless strong evidence exists to the contrary.
In my own work on insect evolution, therefore, I use the follow- ing terminology for wing veins: costa ( + ), subcosta (- ), radius ( + ), radial sector (- ), anterior media ( + ), posterior media ( -) anterior cubitus ( + ), posterior cubitus ( -), and anals ( +, -, or flat). The term postcubitus was suggested by Snodgrass
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19661 Carpenter - Protorthoptera and Orthoptera 51 for the first anal of Comstock and Needham; however, I see no reason to make this change especially since the new name would al- most certainly be confused with Lameere's posterior cubitus men- tioned above.
Order Protorthoptera
As noted above, the Palaeozoic orthopteroids present unusual prob- lems in classification. The Blattodea, although part of this phylo- genetic complex, are not included in the present discussion, since they are usually regarded as comprising a distinct order. The Manteodea and Phasmatodea are as yet unknown in Palaeozoic strata. We are therefore concerned in this discussion with the living order Orthop- tera (i.e., Saltatoria) and with a bewildering variety of orthopteroid fossils, some of which appear to be close to the Orthoptera, but others which are suggestive of the Blattodea, Manteodea, Phasmatodea, Plecoptera, or combinations of two 01- more of these groups. Un- fortunately, our knowledge of about four-fifths of these species is restricted to the fore wings or even to only a part of the fore wings. Handlirsch (1906) recognized two main extinct orders in the complex, the Protorthoptera and Protoblattoidea, but found it neces- sary to recognize a third category, "Protorthoptera vel Protoblat- toidea" for the species which he could not clearly assign to one or the other. As more Palaeozoic insects became known, a gradual diminution of the distinctions between the Protorthoptera and Pro- toblattoidea resulted and the number of genera in the "Protorthoptera vel Protoblattoidea" category became nearly as great as the number in the Protoblattoidea itself. In 1937, Martynov suggested the sep- aration of the several non-saltatorial families into a distinct order, Paraplecoptera, leaving in the Protoi-thoptera only the saltatorial forms. More recently, this proposal has been amplified and somewhat altered by Sharov, who has suggested additional differences between the Protorthoptera, Protoblattoidea, and Paraplecoptera. This in- volves the transfer of a few species (Oedischiidae) with well de- veloped jumping hind legs into the true Orthoptera, restricting the Protorthoptera to one family, having an incipient saltatorial modifica- tion of the legs, with the bulk of the Palaeozoic orthopteroid families going into the Paraplecoptera and Protoblattoidea. Before considering Sharov's proposed classification, I wish to dis- cuss certain aspects of the venation of the fore wing of these Orthop- tei-oids, at least those features which involve differences in interpretation. Sc, RI, Rs, CUP and the anals present no difficulties in their homologies, but the media (and to some extent CuA) is a
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52 Psyche
[March
different matter. In the orthopteroids, as noted above, the media does not show the clear division into a convex anterior branch and a concave posterior one. It is often deeply forked and the posterior branch may be strongly concave or only slightly concave or even neutral (flat), but I think it can be safely said that there is no wthopteroid known in which the anterior branch of the media is convex. We have no way of knowing, therefore, whether in such cases the entire media consists only of MP (with a flattened anterior branch) or of LIA and MP, with a flattened MA. The only positive criterion by which we can identify a vein in the orthopteroids as homologous with MA of the Palaeoptera is by its convexity -which none have. I think there is enough evidence, however, to justify the probable determination of the anterior branch of M as MA in some families of orthopteroids, but the determination is only a working hyp~thesis.~
Another area of controversy is the relationship between CuA and M. In the majority of the orthopteroids there is some type of con- nection between M and CuA if only a short cross-vein. In others (as Stereopteridae, figures 10-13 of the present paper), CuA curves upwards and fuses with part of M before diverging off as an in- dependent vein. It should be noted that there is marked individual variation in the nature and amount of this coalescence. In others. such as the Blattinopsidae, there is a strongly convex stem of M (see figures 7 and 8 of this paper) which become abruptly flat or concave after the divergence of a short, convex, posterior branch. I think it probable here that the anterior branch of CuA is fused with M from the very base until the point of divergence. A somewhat similar situation appears to occur in the Oedischiidae and related families (these being treated here as true Orthoptera), but I believe rhe homologies are different (see figure 15). The stem of M, in- stead of being markedly convex, is flat or even concave. The short vein which diverges towards CuA is rather weak in the Oedischiidae, althoug,h it may be stronger in other, related families. In this case, 'In my own desciiptive accounts of the Paleozoic orthopteroids I use the designation MA and MP if the posterior branch is definitely concave and the anterior branch flat; if the posterior branch is flat like the anterior one I use the designation M for the entire system; if all branches of the media are concave, I use the designation MP for all. EXPLANATION OF PLATE 4
Lmatofihora typa Sellards.
Photograph of specimen No. 3539, Museum
of Comparative Zoology, showing prothoracic lobes, with hair covering and reticulated pattern, Original.
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54 Psyche [March
I consider the divergent vein as a modified cross vein, which in many cf the orthopteroids appears in diverse forms (e.g., Strephocladidae, figures I and 2). It is my opinion, therefore, that the connections between CuA and M are of a diverse nature in the orthopteroids and that these connections have arisen independently many times. Regarding Sharov's proposed classification of the Palaeozoic orthop- teroids, I have previously ( 1954) adopted Zeuner's suggestion (also accepted by Sharov), that the Oedischiidae are true Orthoptera; Sharov has with good reason made a similar inclusion of a few related families (of which the Permelcanidae, figure 18, is a rep- resentative). He then proposed restricting the Protorthoptera to the single family Sthenaropodidae, defining (1960, p. 295) the order as including those orthopteroids with "dorso-ventral flattening of the body, cursorial hind legs, lacking the two rows of spines on the hind margin of the tibia, by the small precostal area lacking the numerous veinlets and by the absence of an undifferentiated concave MA2." This definition I find much too narrow for an order; it might well fit a family - a small one - but certainly not an order. The re- mainder of the oithopteroids which I have previously included in the Pi-otorthoptera, Sharov proposes to divide into the Protoblattodea and the Paraplecoptera. The former order he would restrict to those species having wide coriaceous fore wings, the absence of a clearly defined division of the media stem into two main branches, MA and ^\IF, by large coxae and by general resemblance to Blattodea. In this case, Sharov's characterization seems to be much too broad and generalized. Certainly the coriaceous nature of the fore wings varies greatly within orders (e.g., Orthopteraj ; in some the fore wings are truly men~branous but in others they are definite tegmina or even dytra.
So far as the division of the media into MA and MP is concerned, I question that this is clearly divided in any of the orthop- teroids; as noted above, there is no orthopteroid that has a convex, and therefore, definite, MA. The coxae are known in very few of the species that Sharov would place in the Protoblattodea and, once again, I cannot see this as an ordinal characteristic. The Paraplecop- tera are distinguished by Sharov by the presence of membranous, elongated fore wings, by the clearly defined division of the median into MA and MP and by the general resemblance of the insects to the Plecoptera. On examining the genera which Sharov includes in the Paraplecoptera, as described and figured in the Osnovy (1962 j, I find many families (e.g., Spanioderidae, Probnidae, Strephocladidae, etc.) in which the fore wings are distinctly coriaceous and as rela-
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19661 Carpenter - Protorthoptera and Orthoptera 55 tively broad and oval as those of the previous order. The condition of MA and MP has already been commented upon. I can see no justification in Sharov's account for the recognition of the Protorthoptera, Protoblattodea and Paraplecoptera as separate orders, and I propose to place all of these without subgrouping in the order Protorthoptera. Admittedly, the Protorthoptera as thus con- stituted would bs almost: certainly polyphyletic. But it seems to me that Sharov's classification would recognize two polyphyletic orders, (Pi-otoblattodea and Paraplecoptera) with the order Protorthoptera itsel-f so narrowly defined as to include only one family. In all probability, the Palaeozoic Orthopteroids were not evolving just in che direction of the living orders Blattodea, Plecoptera, and Orthop- tera but, as a result of radial evolution, in many directions. Certainly this is what one would expect from the geological record of other groups of animals. The setting up of the three orders Protoblattodea, Paraplecoptera and Protorthoptera would seem to me to conceal what were almost certainly the real evolutionary lines of these insects. Hence, I prefer to group these orthopteroids into one large complex t h e Protorthoptera- until we have enough evidence to indicate what the several lines of evolution have been. I do not believe that we have that now.
I am convinced that Sharov is correct in maintaining that the Lem- matophoridae are not sufficiently different from the Liomopteridae, etc., to justify separation in a distinct order, Protoperlaria. Certainly, as Sharov points out, both fore and hind wings of the Lemmato- phoridae and related families can be distinguished from those of other Protorthoptera only with the greatest difficulty. I cannot agree wtih Sharov! however, in his claim that the paranotal lobes in the Lemmat~pho~idae were continuous and formed a pronotal shield as in Liomopteridae, instead of being independent lobes, as Tillyard and I had described them. Sharov states that his study of the pub- lished photographs in till yard'^ ( I 928) and Carpenter's ( I 935 ) papers shows that the lobes unite in front and behind. Although
photographs are extremely useful in the study of fossils, they are no substitute for the actual specimens.
Till yard's drawing and mine
were based on different specimens and were made several years apart. I have re-examined the material in both the Harvard and Yale col- lections since the publication of Sharov's paper and I cannot agree with the interpretation which he has made from the published photo- graphs. Photographs of the thoracic region of two specimens of Lemmatophora typa Sellards are included here (plates 4 and 5). The first of these shows a specimen which is not quite in a symmetrical
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position; it shows especially clearly the form of the individual lobes. The second specimen, which is the one originally figured by Tillyard, shows the thorax in a more symmetrical position. When Tillyard's original photograph was made. plant fragments and other organic debris covered much of the thorax, obscuring the form of the para- notal lobes posteriorly. Subsequently, as shown in the photograph on plate 5, this debris was removed, presumably by Tillyard himself. The paranotal lobes are reddish-brown in color, like the true wings; the plant fragments and the debris are black, so that the two are more distinctive in the actual fossil than is apparent in a black-and- white photograph. In any event, I do agree that these paranota are not sufficient to justify the separation of the Lemmatophoridae from the Protorthoptera.
In the preceding papers in this series, eight families of Protorthop- tera were considered: Lemmatophoridae, Probnidae, Liomopteridae, Chelopteridae, Stereopteridae, Demopteridae, Phenopteridae and Protembiidae. In the present paper three additional families are covered, the Strephocladidae, Blattinopsidae, and Tococladidae, and the Stereopteridae are discussed further, in the light of new material. Family Strephocladidae Martynov
Strephocladidae Martynov, 1938, p. 100.
Fore wing: coriaceous ; precostal area absent; Sc well developed, extending to mid-wins; or beyond, with several to many forked branches; Rs arising before mid-wing; RI extending well towards iipex, with several oblique branches leading to margin beyond Sc; Rs very well developed, with several to many long branches, usually without forks except for the branches in the apical part of the wing; M forked before origin of Rs, the anterior branch often touching Rs briefly or connected to it by a short, stout cross vein; M with sev- eral long branches, usually simple, independent of R basally, often touching CuA briefly or connected to it by a stout cross vein or pos- sibly by an anastornosed branch; Cu independent of M basally; CUP arising near base ; CuA directed longitudinally, giving rise to several long branches, usually simple; branches of Rs, M and CuA parallel and slightly sigrnoidal; CUP usually nearly straight, except near its distal end; a distinct furrow posterior to CuA, very close and parallel to it; IA close and parallel to CUP; other anal veins irregular and EXPLANATION OF PLATE 5
Lmatophora typa Sellards, Photograph of specimen No. 5115, Peabody Museum, Yale University.
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58 Psyche [March
higlily variable; cross veins numerous and regularly arranged over die areas of Rs, M and CuA; an irregular network in costal area 2nd between CuA and CUP and the anal veins. Wing membrane with fine microtrichia between veins; prominent setae or other cuti- cular d~rivatives developed to variable degrees on most of the veins of wing. including many cross veins.
Hind wing unknown and body structure unknown, except for part of one leg.
This family has not been given any diagnosis previously, except by generic assignment. In the preceding account I have attempted to bring together venational characteristics of the fore wing present in several Palaeozoic genera which are apparently closely related to Sti-ephoclfidus and which I am placing in the family. However, the sti-ephocladids will probably turn out to be an extensive group and its diagnosis will undoubtedly need modification as other genera be- come known.
S/r(>phocladiis was established in the Order Palaeodictyoptera by Scudder (1885) for a species (subtilis Kliver) which was collected in Uppei- Carboniferous strata of Saasbrucken and which had osig- inally been placed by Kliver in the blattod genus Petroblattina, It
has subsequently been placed in the order Protoblattoidea, Incestae Sedis, by Handlirsch
( 1908, 1921 ) ; in the order Protorthoptera, family Oedi~chiiclae, by Waterlot ( 1934) and Guthorl ( 1936) ; in a new order Strephocladodea, family (new) Strephocladidae, by Mar- tynov ( 1938) ; ii-i the order Paraplecoptera, family Stsephocladidae by Sharov ( 1961) ; and in the Protosthoptera, family Strephocladidae, bj Kilkaio~ 5 ( 1965), who added another genus, Spargoptilon (L. Permidn, Aloravia), to the family. From my study of the type speci- men of Strcphocldus subtilis and of the several species from Elmo CUP
Text-figure 1. Strephocladus subtilis Kliver. Original drawing of fore wing based on holctype, No. D/164, Bergingenieurschule, Saarbriicken.
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19661 Carpenter -- Protorthoptera and Orthoptera 59 described below, I consider that the family Strephocladidae fits well within the order Protorthoptera and that Martynov's order Strepho- cladodea is synonymous with the order Protorthoptera. The relation- ships of the family Strephocladidae within the Protorthoptera are not so definite. The little-known Strephoneuridae Martynov, from the Lower Permian of URSS, are closest in venational details so far as they are known; but only when the hind wings and body struc- tures have been found can these affinities be worked out satisfactorily. The most significant features of the Strephocladidae are the pres- ence of long, forked branches on Sc, and the long, parallel and nearly unbranched veins forming Rs, M, and CuA. The identity of most of the main veins is clear; RI and CuA are strongly convex, and Sc, RI and CUP are concave. The media does, as usual in the Protorthop- tera, present a problem; it shows neither convex nor concave elements and is accordingly being designated here as M. In Strephocladus, Spargoptilon and the new genera herein described, CuA either arches anterior, touching M briefly (Spargoptilon) or connects with M 'by a stout- cross vein; because of the convexity of all veins included, I agree entirely with Dr. 1Cukalov~'s interpretation that no branches of M are, in fact, involved in the CuA complex. The relationship between Rs and the anterior branches of M seems to be similar; in some species (Spargoptilon) there is slight anastomosis, but in others the connection is bv a cross vein. These variations almost certainly occur as individual fluctuations within species. Apart from the general venational pattern, there are two features of the fore wings of strephocladids that deserve further comment. I). Setae on veins. Most orthopteroids possess fine microtrichia on ;he wing membrane and their presence on the wings of Protorthop- tera is well known. The notable feature here is the presence of large setae on the veins, these being especially clear in å´Homocladus These are, of course, represented in the main by setal bases, the setae them- selves apparently being broken off in the rock matrix. These setae occur only on that half of the fossil (reverse) which has the impres- sion of the dorsal surface of the wing; the ventral surface of the wing was apparently devoid of such setae. Setae have previously been found on the veins of a few Protorthoptera but they have not previously been noted as occurring so abundantly or regularly. Neither microtrichia nor setae are visible on the type specimen of Strepho- cladus subtilis but this is almost certainly the result of poor preserva- tion of the fossil. (2) The costa, for a variable distance along the anterior margin, is actually sub-rnarginal for most of its length, there being a narrow but distinct, membranous border. This is a con-
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60 Psyche [March
dition which occurs in many Osthoptera. Under low power magni- scation, this gives the impression that the costa is a much wider vein than it actually is. (See Plate 6).
Since the figures and descriptions of Strephocladus subtills Kliver which have previously been published are not satisfactory, I include here (Text-figure I) an original drawing of the type, which was placed at my disposal by Dr. G. Kneuper. The length of the pre- served part of the wing is 20 mm.; comparison with other Strepho- cladid wings indicates that the complete wing was about 30 mm. There are three distinctive features of the wing that separate Strepho- cladus from other genera now known in the family: the presence of short, oblique veins from Sc to the costal margin, the definite termina- tion of Sc on RI ; and the pectinate origin of the branches of CuA. As can be seen in the figure, the front branch of M is in brief contact with R; CuA is joined by a short cross vein to M. It should be noted that the stem of M is not convex, and that the vein designated CuA is entirely convex. The shallow furrow, directly posterior to CUP, can clearly be seen.
The following are the strephocladids in the Elmo limestone: Genus Homocladus, new genus
Fore wing: costal margin with a distinctly arched border at about the level of the origin of Rs; Sc with numerous long, branched vein- lets, directed longitudinally and terminating on costal margin; Rs arising at about one-third the wing-length from the base, giving rise to numerous, long branches, all simple except near the wing apex; M forked just before the origin of Rs, forming several long, simple branches; fork of CnA at least slightly basal of the first fork of M; CUP at its distal end extending parallel and close to the wing mar- gin, this marginal vein being continuous basally by extensions of the R s
CUP \ CuA
Text-figure 2,
Homocladus grandis, n. sp. Drawing of fore wing (holo- type)
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19661 Carpenter - Protorthopiera and Orthopirrd 61 anal veins; branches of CuA arising di~hotomously, more cumed than tl~ose of Rs and A1 ; apex of wing much neal-er the anterior wing margin than the posterior. Area between CuA and CUP with a coarse retidation of cross veins, Hind wing unknown. Type-species: Iiotnocl~dus grandis, n. sp. , -
1 his genus differs from &rtphocIadw in lacking the straight, oblique branches to the costa, and in having the branches of CIA arising dichoton~ousIy, instead of pectinafely ; it differs from 8pargo$- MOQ by its more slender wing shape, the pxtinate, instead of dichot- omous, branching of Rs, and in the position of the distally extended part of CUP,
Homocladus grandis, n, sp.
Text-figures 2, 3, 4 and Plate 6
Length of fore wing (holotype) : 43 mm.; width, 11 mm. Cmtal area with only a slight broadening before mid-wing; several rows of cells between CuA and CUP; wing without markings; setae Text-figure 3, Hornoddu~ prfiniis, n. sp. Photograph of wing surface of hdatype, showing veins (1') and setal bases (M) on yein3 and membrane; and mkrotrichia on membrane.
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62 Psyche [March
numerous and well developed on main veins and some cross veins. The venational details of the holotype are shown in figure 2. Holotype: No. 58742b, Museum of Comparative Zoology; col- lected by F. hf. Carpenter) in the lower layer of the Elmo limestone in 1927. The specimen consists of a very nearly complete fore wing) jacking only the distal wing margin. A second specimen (No. MCZ 5875ab), with the same collecting data, consists of the proximal two- chirds of a fore wing; a drawing is included here to show the ap- parent fluctuation in the venation. Also on this piece of rock, only 2 or 3 mm. from the wing, is part of a femur and tibia of a leg; the proximity and size of this leg indicate that it is from the same insect as the wing. The tibia is armed with two rows of heavy spines and the femur bears a few smaller ones.
Homodadus ornatus, n. sp.
Text-figure 5
Fore wing: Iefigth, as preserved, 20 mm. ; width, 7 mm. ; estimated complete wing length, 30 mm. Costal area with a more prominent broadenhg than in grandif; area between CuA and CUP with fewer cells; wing at least four transverse bands. Venational details are
shown in figure 5.
Holotype: No. 15584, Peabody Museum) Yale University; col- lected in Elmo limestone by C. 0. Dunbar) 1921. This species difiers from grandis mainly by the wing markings 2nd smaller size.
Genus Paracladus, new genus
Fore wing: costa1 margin almost smoothly curved; Sc with sev- eral oblique veinlets, mostly branched, but generally much less de- vel~ped than in FIo~tzocZud~~s; Sc apparently terminating either on Text-figure 4.
Humucladus grandis, n, sp. Drawing of fore wing (para- type no. 5875ab).
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19661 Carfienter - Protorthofitera and Orthofitera 6 3 margin or RI ; origin of Rs at or slightly beyond mid-wing, its branches arising dichotomously; CUP only very slightly curved dis- tally, without the terminal extension as in FIomocZadus. bIicrotricl~ia on wicg membrane more numerous than in HonzocZadus; setae on veins apparently less developed and less numerous. Type-species : PmacZadm retardatzis, n. sp. This genus, though clearly related to &IonzocZadus, differs by the late origin of Rs, and especially by the absence of the rna~-ginal ex- tension of CUP; in the latter respect) it resembles Strephoclailus. Paracladus retardatus, n. sp.
Text-figure 6
Length of fore wing, as preserved, 20 mm.; width, 7 mm.; esti- mated complete wing length, 30 mm. Subcosta space distinct and rather wide, with a few cross veins; relatively few cells between CuA and CUP; IA connected to CUP distally by a strong, longitu- dinal cross vein. Wing markings absent. Venational details a]-2 shown in figure 6.
HoIotype: No, 5877ab, Museum of Comparative Zoologjr; col- lected in the lower layer of Elmo limestone in 1932 by J?. 31. Cas- penter.
Family Blattinopsidae Bolton
Blattinopsidae Bolton, 1925, p. 23.
Oryctoblattinidae Handlirsch, 1906, p. 705 (OquYobl~ttina is a jun!or objective synonym of Blattinop~is) .
Fore wing : mcmbsane apparently thin) at most weakly coria::~:; ; wing generally oval but often very short; Sc extending at leasf ::) mid-whg, often conside~ably beyond; area between Sc and RI at IF:.:; Text-figure 5, kiomocladz~s ornm'us, n. sp. Drawing of fore wing (holo- type) -
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64 Psyche [March
as wide as the costal space; RI usually slightly sigmoidal; Rs usually arising at about one-quarter of the wing length from RI and giving i-ise to several branches either dichotomously or pectinately; M ap- parently independent of R at the base) although very close to it in some species; M almost always with at least two terminal branches; CUAI apparently anastomosed with the stem of M for a considerable distance) divei-gi~~g posteriorly from it just before the level of the origin of Rs, then coalescing for a variable distance with CuA2; CUP arising from die stem of Cu near the base of the wing and continuing as a nearly straight> strongly concave vein ; the posterior margin of the wing is usually strongly indented at the termination of CuF; IA usually very close to CUP and in some species apparently xnastomosed with it; usually at least two other distinct anal veins present; cross veins highly variable in form) usually numerous and often forming a reticulation over the central and posterior portions of the wing; a cwved transverse line) starting from RI at about the level of the end of Sc and terminating on CuA at about the level of :he first definite fork of CuA) is visible on the wings of most mem- bers of the family.
The hind wings and body of the blattinopsids are almost entirely unknown; a short, stout ovipositor apparently existed in some species (Kukalovi) 1959). A fragmentary wing) probably a hind wing) is the basis of the descripticn of Blattinopsis eleyans Handlirsch ( 1906, p. 160)~ from the Upper Carboniferous of Germany; however, since this wing is not associated with a fore wing and since the venation of the re~nigium is distinctly different from that of the fore wing) there is no real basis for considering this to be a member of BZat- tino$sis. Laurentiaux ( 1950, p. 66) has established Blattinopsis in- certa (Upper Carboniferous of France) for a specimen consisting of Text-figure 6.
Paracladus retardatus, n. sp. Drawing of fore wing (holo- type).
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19661 Carpenter - Protorthoptera and Orthoptera 65 2 fore wing and part of a hind; however, venational details of the fore wing) such as the proximity of Sc to RI, eliminate this species from Blattinopsis; it is herein assigned to the genus Stephanopsis KukaIovi, which was erected as a subgenus of Blattino~sis in 1958 ( p. I 3 1 ) , with incerta as the type species of the subgenus. This is the only species, apart from the very dubious elegans Handlirsch, previously mentioned? which can be assigned at the present time to Stephanopsis. It is highly doubtful, in my opinion? that Stephanopsis actually belongs to the Blattinopsidae.
The venation of the fore wings of the blattinopsids presents some difficulties) at least with respect to homologies of M and CuA. In all members of the family which have been described? the basal
portion of M is strongly convex and the remainder concave or at leas: neutral. The change in the topography of this veins occurs as 2, strongly convex branch diverges obliquely, fusing with what is obviously part of the anterior cubitus, as shown in figure 7 and in the numerous illustrations of blattinopsids given by Kukalovii (1959, 1965). The venation oi the blattinopsids, in this respect, is different from that of the carcurgids (i.e., ~Heterologus) and the oedischiids in having no concave vein between CLIP and CuA. It seems most likely to me that CUAI is coalesced with M basally and that it then diverges off as the oblique vein and anastomoses with CuAz. Sharov (1962) is of the opinion that NIP is the oblique vein that coalesces with CuA. This interpretation? however, does not explain the strong convexity of the base 01 M, the convexity of the oblique vein itself, or the change in the topography of the rest of M beyond the diver- gence of the oblique vein.
In some genera of blattinopsids ( i.e.) G~aphyrophlebia) distinct grooves extend longitudinally between the branches of Rs and M. They have been represented in some figures (Handlirsch? 1906) as actual veins but examination of these wings under high magnification and optimum illumination fails to show any sign of cuticular lines along the grooves. Actually, these structures seem to be shallow depressions in the membrane bordered by low ridges of membrane; similar surface features are found in the wings of various genera of orthopteroid insects? including the Blattodea. The most notable structure in the blattinopsid fore wings is a curved line which runs transversely from RI at about the middle of the wing to CuA or even slightly beyond. Kukalov6 ( 1959) has
pointed out that this resembles the line in the fore wings of some Recent cockroaches of the family Polyphagidae, in which it is ap- parently formed by spreading and folding of the wings. Others have
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[March
noted the similarity of this line to the transverse mark that occurs in the fore wings of some Homoptera, and Haupt (1941, p. 88) has actually established a new order, Protofulgorida, for the Blattinop- sidae, which he regards as closely related and ancestral to the Homoptera. All available evidence, however, indicates that the blat- tinopsids are undoubtedly orthopteroid. For example, the details of venation are surely like those in other Protorthoptera and the anal area is clearly orthopteroid, not homopterous. Previous accounts of the blattinopsids (Bolton, 1925 Kukalovi, 1959) have noted the fragmentary nature of all specimens of fore wings. So far as I am aware only one species, Glaphyrophlebia spe- ciosa (Sellards), is known from a complete wing. In most specimens) either the apical region or the anal area has been broken away. This is true even of such relatively large species as Blattinopsis kukalovae, described below, and it is in marked contrast to the frequency of occurrence of undamaged wings of such small and delicate insects as the Homoptera and Psocoptera in the Elmo limestone. This cir- cumstance seems to indicate that the fore wings of the blattinopsids were unusually thin and delicate.
Genus B lattinopsix Giebel
Blattinu$~sis Giebel, 1867, Zeitschr. Ges. Naturw. 30:417; Kukalovi, 1959, Rozpravy. Ceskos. Acad. Ved. 69 : (1) :5. Oryctublattina Scudder, 1895, Bull. U.S. Geol. Surv. 124:133 (jr. obj. syn.). Fore wing: costal area and area between Sc and RI with numerous oblique veinlets, very close together and often branched; area between branches of Rs, M and Cu with similar cross veins, those in the Text-figure 7. Blattinu#sis kukalovae, n. sp. Drawing of fore wing (holotype). M+CUAl is the convex base of these coalesced veins; T is the transverse line.
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68 Psyche [March
distal part forming a fine reticulation; area between CuA and CUP with numerous oblique, parallel branches, mostly slanted towards the posterior margin and parallel to CUP; anal area with numerous cross veins forming a reticulation.
Generic limits of Blattinopsis are very difficult to make because of the obvious variation in the venation within species. I consider it advisable to treat this genus broadly, as was done by Kukalovi. The type-species, Blattina reticulata Giebel, is not so well known as several others, but there is no question about its basic venation. The genus has been recorded from Upper Carboniferous and Lower Per- mian strata of both Europe and North America. The probable generic synonomv has been discussed by Kukalovi ( I 965 ) . Blattinopsis kukalovae, n. sp.
Text-figure 7
Fore wing : incompletely known ; estimated length, 22 mm. ; hslo- type
(basal half of wing), 12 mm.; width 7 mm. Costal margin nearly straight, not conspicuously arched; costal area narrowed at wing base; venation typical of the genus; Rs arising from a single stem; M (in holotype and ~arat~pes) with a deep fork; cross vein;; in costal area and between RI and Sc, and RI and Rs very close together and parallel, almost without cellules; reticulation between branches of Rs, M and CuA.
Holotype: No. 6301ab, Museum of Comparative Zoology; con- sisting of basal half of a wing; collected by F. M. Carpenter in the upper layer of the Elmo limestone.
Paratype: No. 6302ab, Museum of Comparative Zoology; con- sisting of a more distal portion of the wing, lacking the apex; para- type No. I 5 582ab, Peabody Museum, Yale University ; collected by C. 0. Dunbar; middle portion of the wing, not so well preserved. A third specimen, No. 15633b, Peabody Museum, is a small frag- ment which probably belongs to this species. This species has the wing form and size of the type-species of the genus, but it lacks the reticulation in the area between RI and Sc, and the subcosta is considerably shorter than in reticulata. It differs from most of the species described by Kukalovh from the Permian of Czechoslovakia in having (I) the costal margin smoothly curved, (2)
less reticulation of the cross veins and (3) a much less con-
spicuous lobation of the anal area.
The species is named for Dr. Jarmila Kukalovh of Charles Uni- versity in Prague, in recognition of her achievements in both collect- ing and studying the Permian insects of Moravia.
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19661
carpenter^ Pro forthoptera and Orthoptera Text-figure 8. Glaphyrophlebia speciosa (Sellards) . Drawing ef fore wing, based on neotype.
Genus Glaphyrophlehia Handlirsch
Glafhyrophlebia Handlirsch, 1906, Proc. U.S.N.M. 29907. Sindon Sellards, 1909, Amer. Journ. Sci., 27 :I%. h a Sellards, 1909, ibid. 27:153.
Fore wing: membranous or weakly tegminous; Sc terminating at or slightly beyond mid-wing; costal and subcostal areas with few cross veins; Rs arising at or slightly before mid-wing, with numerous branches, most of them forked distally; M and Cu as in BIat/inopsis, but usually with fewer branches leading from CuA to the hind mar- gin distally; areas of Rs and M with a few, widely scattered distinct cross veins, not forming a reticulation; curved line across middle of wing, much as in Blattin0i)sis; space between branches of Rs and M with a prominent groove or grooves parallel to the veins. Hind
wing unknown.
Type-species : Ghph yrophlebia pusilla Handlirsch from Grove County, Pennsylvania, Illinois; Upper Carboniferous. The type- specimen of pusilla consisted of the distal two-thirds or half of a fore wing;6 although it is well preserved, Handlirsch did not dis- tinguish between the actual forks of the branches of Rs and the grooves in the intervening membranes. Almost certainly the veins had two distal forks, as in the other species now known in the genus. "The type and only known specimen of fusilla was contained in the Daniels Collection at the time of Handlirsch's description. This collection was sup- posedly turned over to the U.S. National Museum (Handlirsch, 1906, p. 662) but apparently it was not; none of the specimens originally in the Daniels Collection are in the National Museum.
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70 Psyche [March
This genus is undoubtedly close to Blattinopsis but differs from it by having (I) distinctly few cross veins over the wing in general and especially in the distal portion and (2) clearly defined grooves in the wing membrane between the branches of Rs and M. Sindon Sellards and Piirsa Sellards seem to me to be inseparable from Glaphyrophlebiu,: as shown below, the type-species of both of these genera possess the venational characteristics of Handlirsch's genus. Glaphyrophlebia is represented by one species (clava Kukalovi) from the Lower Permian of Moravia and two species (uralensis Martynov and rossicz~m Martynov) from the Permian of USSR, both orig- inally described in Sindon.
Glat>lzyrophlebia speciosa (Sellards)
Text-figure 8
Sindon spec'iosa Sellards, 1909, Amer. Journ. Sci., 23 :154, fig. 1. Fore wing: length, 8 mm., width, 3.8 mm. (neotype). Sc ter- minating at mid-wing; RI strongly sigmoidal; Rs with seven main branches, each forked distally; M (in type, probably variable) forked only" near wing margin; oblique part of CuA slightly basal to the origin of Rs; CuA with only about six branches leading to the hind margin ; costal veinlets unbranched (in type), separated by spaces about equal to their length; cross veins in the area of the subcosta and Ri with similar spacing; veinlets from RI to the costal margin beyond the end of Sc somewhat more numerous and closer together; cross veins in the area of Rs, M and CuA widely spaced; no reticula- tion between CuA and CUP, although two rows of cells occur in that area basally; anal area without a reticulation. The holotype specimen, No. 85 in the Sellards collection, was studied by me in 1927, at Austin, Texas. Since this fossil has sub- sequently been lost, T designate as the neotype specimen No. 6303, in the Museum of Comparative Zoology. This was collected at Elmo, by F. M. Carpenter in 1927; it consists of a complete and well pre- served fore wing. Sellards figure of the original type was slightly in error in showing Sc too long and in showing too many branches from CuA to the wing margin. The neotype is very close to the original type except that in the latter the fork of M was much deeper. As pointed out by Kukalovii ( 1965), the branching of M is subject to much fluctuation within the species of Blattinopsidae. ? -^
1 he venationai details of this species are shown in text-figure 8. So far as I am aware, this is the only species of the Blattinopsidae known from a complete wing.
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19661
Carpenter - Protorthofitera and Orthofitera Text-figure 9. Glaphyrophlebia ovata (Sellards). Drawing based on neotype.
Glaphyrophlebia ovata (Sellards)
Text-figure 9
Pursa ovata Sellards, 1909, Amer. Journ. Sci., 27:156, fis. 4. Fore wing:
length 8 mm.; width, 3.5 mm. (neot~pe). Sc ter- minating at mid-wing ; RI slightly sigmoidal; Rs with 9 branches, each forked distally (neotype) ; M forked only at wing margin (neo- type) ; costal veinlets and those between RI and Sc much closer together than in speciosa; cross veins between RI and anterior mar- gin numerous and close together; cross veins between RI and Rs and between branches of Rs much as in speciosa; CuA with about 13 branches leading to hind margin, close together and parallel; area between CuA and CUP with a reticulation basally. The holotype specimen, No. I 126 in the Sellards collection was studied by me in 1927; since tha-t has subsequently been lost, I desig- nato as the neotype specimen No. 4965ab, Museum of Comparative Zoology; this was collected by F. M. Carpenter, at Elmo, in 1927. It consists of a very well preserved fore wing, lacking the anal area. ,?
I he species is similar to sfieciosa but has the veinlets in the costal and subcostal areas much more numerous; it also has more branches of Rs and of CuA. The grooves of the membrane between the branches of Rs and M are more complicated than in speciosa, each one apparently being composed of 2 or 3 fine grooves. Sellards figure of ovata, based on a poorly preserved wing, con- fused the branches of Rs with grooves between them and also incor- rectly represented the structure of M. This species has the relatively small number of cross veins and lack of reticulation characteristic of Glyphrophlebia but has more cross veins and veinlets from CuA to the hind margin than speciosa does; in these respects it suggests the condition in Blattinopsis more than the latter.
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Text-figure 10. Stereopierum breve, n. sp. Drawing based on holotype. Family Stereopteridae Carpenter
Stereopteridae Carpenter, 1950, p. 201.
This family was established on a single species, Stereopterum rotundurn Carpenter, from the Elmo limestone. Several additional specimens of this insect have subsequently been found as well as representatives of two other species, described below. In one of the latter (S. breve, n. sp.) the prothorax is seen to be broad, with the pronoturn extended laterally, but truncate anteriorly and posteriorly. Nothing further is known of the body structure in this family. The stereopterids were probably related to the Euryptilontidae, known from the Lower Permian of the USSR. The prothorax seems to be quite differently formed in these two groups and until the hind wings are known, it seems advisable to recognize the families as dis- tinct.
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19661 Carpenter - Protorthoptera and Orthoptera 7 3 Stereoptermn rotund~um Carpenter
Text-figures 1 I, 12
Stereopterum rotundurn Carpenter, 1950 Proc. Amer. Acad. Arts Sci. 78: 202.
Six additional specimens have been collected in the Elmo lime- stone since 1950, these bearing numbers 4959, 5253, 5254, 5258, 5259, 5260; all were collected in the lower layer of the limestone (F.M.C.).
The species has not yet been recorded from the upper layer.
A survey of all of these fossils shows a greater variation in the venational pattern of the fore wing than has previously been realized.
The subcosta and RI remain fairly constant. Rs in sev- eral specimens (4959, 5880, 5885) has a deep fork; in another (5887) it has three branches instead of 2 and in a third (5879) it is unbranched. In the type specimen (4922) it has a shallow, distal fork. In a few specimens (5886, 5879, 5885) MA1 is anastornosed for a short distance with Rs; in others, as well as the type, it is free from Rs. The weak condition of the basal part of CuA2, as seen in the type, is apparently unusual; in most specimens (e.g., 5887, 5879) it arises distinctly from the stem of CuA; in one specimen (5886) it arises from CuA+ M, i.e., before the separation of CuA from M. The pattern of cross veins is variable but essentially as shown in the type; beyond the end of Sc the oblique veinlets from RI to the margin are somewhat closer together than elsewhere; this seems to be consistent in all specimens. The distinctive cluster of hairs on MA and MP near the middle of the wing can be seen in several specimens in addition to the type (e.g., 5879) ; few smaller setae are visible on some other veins but they do not form a definite patch.
All of the fossils which show the base of the wing have a distinct lobe which resembles a heavily sclerotized fold of the costal area extending backwards of the base of Sc and R (see text-figure 11). Having noted this in several specimens, I attempted to chip away the very base of the type specimen, which turns out to have the lobe present also. This lobe was roba ably concerned with the fitting of the tegmen against the pronoturn when the wings were in the resting position; a variety of sclerotized structures, which occur at the wing bases in many living orthopteroids, seem to have a similar function.
Drawings of two specimens are included here (text-figures 10 and 1 1 ) to show the extreme variation in some of the features mentioned. It should be noted that in one of the specimens (5886) there is no
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74 Psyche
[March
basal connection between CUP and CuA although the latter pro- duces both CUAI and CuA2 directly from M. In the other specimen (4958) both CUAI and CuAa arise from .MP, not from the stem of M.
sc
CuA
s c
Text-figure 11. Stereopterum rotundum Carpenter. Drawing of speci- men no. 5886, M. C. Z.
Text-figure 12. Stereoptcrum rotundum Carpenter. Drawing based on specimen no. 4958, M. C. 2.
Text-figure 13,
Stereopterum maculosum, n. sp. Drawing based on holo- type.
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19661 Carpenter - Protorthoptera and Orthoptera 7 5 Text-figure 14. Tococladus rallus, n. sp. Drawing of fore wing (holo- type
Stereopterum maculosum, n. sp.
Text-figure I 3
Fore wing: length, 8.5 mm., width, 3.2 mm. (holotype) ; more slender than that of rotundurn; Rs usually with some coalescence between Rs and MA1 ; CuA or CuA1 anastomosed with stem of M; cross veins in area between CuA and CUP forming a very coarse network; wing distinctly coriaceous, with numerous irregular macula- tions, preserved as reddish-brown spots. Holotype : No. 15653, Peabody Museum of Natural History, Yale University; collected in the Elmo limestone by C. 0. Dunbar. This consists of a nearly complete wing, lacking the anal area and portions of the wing base and anterior margin. Paratypes: No. 5257, Museum of Comparative Zoology; collected in the lower layer of the Elmo limestone by F. M. Carpenter; this consists of a nearly complete fore wing, lacking only the basal part; a plant fragment rests near mid-wing, obscuring the veins in a small area. No. 4958, Museum of Comparative Zoology, collected in Elmo limestone, lower layer, by F. M. Carpenter. This consists of a nearly complete fore wing lacking the apical quarter and parts of the hind margin and anal areas. This is the only specimen of maculosum which shows the basal part of the wings; the peculiar lobe at the base of the costal area is not present, although there are several prominent spines along the costal margin basally, much as in rotundum. Apart from its shape, maculations, and more pronounced coriaceous texture, this wing differs from that of rotundum by the more widely spaced cross veins and the coarser network between CuA and CUP. Stereopterum breve, n, sp.
Text-figure I o
Fore wing: length, 7.5 mm.; width, 3.5 mm.; Sc extending nearly to the termination of RI ; only I or 2 veinlets from RI to costal
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76 Psyche [March
margin beyond Sc; costal space forming the basal, posterior lobation as in rotundurn- RI with one or more distal branches; CuA2 absent as a distinct vein; area between CuA and CUP small; cross veins much as in rotzmdunz. Pronotum broad, with curved lateral exten- sions; anterior border concave; with very little extension; no pos- terior extension. Head of moderate size. Holotype: No. 2137, Museum of Comparative Zoology, collected in lower laver of the Elmo limestone by I?. M. Carpenter. This consists of a complete insect, showing both fore wings and parts of the body. Unfortunately, the hind wings are overlapped and rest on the abdomen in such a way that their venation cannot be untangled. This species differs from rotundum and maculosum by its much smaller size, broader fore wing, relatively longer Sc, the apparent loss of CuA2 and the presence of a smaller space between CuA and CUP.
The preservation of both fore wings enables a comparison of the venation of the two wings in this one specimen. As can be seen' in figure 10, there are differences in the branching of all main veins in the two wings, especially of MA, MP, and CuA. In all probability the coalescence of MA with Rs is subject to similar fluctuation, al- though it occurs in both wings. The prothorax is clearly preserved, showing the pronotum proper and the thin, lateral extensions. As shown in the figure, the extensions are absent posteriorly and are scarcely present anteriorly, a1 though they are well developed laterally. This is in contrast to the structure in the liomopterids as well as in the euryptilontids, in both of which the extensions are well developed posteriorly.
*.
I he head is preserved in a dorsal view and shows no structural details, apart from indications of small compound eyes. Family Tococladidae, new family
Fore wing: costal marginal; wing margin very nearly straight, narrowed basally; Sc extending to about mid-wing, with oblique, unbranched veins leading to margin; similar branches from R to margin beyond Sc; Rs arising well before mid-wing, with several long branches; M independent of R basally; MA with long, simple branches; MP unbranched; area between CuA and CUP traversed by numerous, strong cross-veins, not forming a reticulation ; CUP nearly straight, not extending markedly along posterior margin; no separate vena dividens; anal veins numerous and well defined; cross veins distinct and simple over virtually all the wing, including the
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19661 Carpenter~ Protorthoptera and Orthofitera 77 anal area.
Microtrichia and setae absent. Hind wing and body un- known.
r 7
1 his family is probably related to the Ischnoneuridae and Proto- kollaridae. The very base of the wing is known; a precostal area may have been present. The wing is made distinctive by the ex- tensive development of the radius and the parallel arrangement of the branches of Rs and M. The basic structure Cu is different from that of the Oedischiidae and related groups in that all the branches of Rs are markedly convex, without the basal concave vein present in :he Oedischiidae and the Carcurgidae. The stem of M is flat and very weak, not strong and convex as in the Blattinopsidae. Genus Tococladus, new genus
Fore wing: slender, costal area moderately narrow; Sc ending on RI, with numerous oblique veinlets; Rs remote from RI near mid- wing but approaching it distally; anterior branch of M forked shortly after origin of Rs, its branches long and simple; several anal veins or main branches.
Type-species: Tococladus rallus, n. sp.
Tococladus rallus, n. sp.
Text-figure I 4
Fore wing: length, 24 mni.; width, 7 mm. Rs with six branches; front branch of M coalesced with Rs for a short distance before diverging posteriorly (possibly an individual fluctuation) ; IA ap- parently with three long branches; cross veins widely spaced over most of wing.
Holotype: No. 5866013, Museum of Comparative Zoology, collected in the lower layer of the Elmo limestone by I?. M. Carpenter. This consists of a complete fore wing, well preserved but lacking the very base. The wing, which is preserved with fine wrinkles, was apparently thin and membranous, not coriaceous. Order Orthoptera
The Palaeozoic families Oedischiidae, Tcholmanvisiidae and Per- melcanidae are now generally regarded as orthopterous, rather than prororthopterous, this view being based mainly on the saltatorial modification of the hind legs and the probable lateral flattening of the body. Of these, only the Oedischiidae are known from the Upper Carboniferous. These early Orthoptera are not extensively repre-
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78 Psyche [March
Text-figure 15. Oedischia dliamsoni Brongniart. Original drawing, based on type in Inst. Paldont., Paris. pc is the precostal area. sented in any Permian deposit. In the Elmo limestone there is one genvs which appears to belong to the Oedischiidae and which is close to Metoedlschia from the Permian of the Soviet Union. There are also representatives of three other families, two of them new. It should be noted that there are many additional orthopteroids present in the Yale and Harvard collections from the Elmo lime- stone, almost all of them undescribed. But none of them, so far as can be determined now, belong to the Orthoptera. They will be treated in the next part of this series of papers. Family Oedischiidae Handlirsch
Oedischiidae Handlirsch, 1906, p. 700.
Fore wing : usually thin, only slightly coriaceous ; precostal space well developed; Sc extending well beyond mid-wing, with several branches; 31 dividing at about a third of the wing-length from the base into MA, which is not clearly convex, and MP, which is mark- edly concave; MA usually anastomosed with Rs; Cu forked near the base, the posterior branch, CUP, strongly concave; the anterior branch (CuA) forking at least once, in part coalescing with a cross vein leading from M, and eventually forming several branches which lead to the hind margin ; CUP unbranched; at least three anal veins; cross veins numerous, in some species forming a reticulation in certain parts of the wing. (See text-figure I 5) Hind wing unknown, Body unknown, except for legs; hind legs long and modified for jumping.
This family is known from the Upper Carboniferous of France (Cornmentry) and Germany and the Permian of USSR and pos- sibly Czechoslovakia. The species described below constitutes the first record of the family in North American deposits. Unfortunately, almost all specimens of Oedischiids consist of iso- lated wings. The type-species of Oedischia (williarnsoni Brong- niart) is represented by two fragmentary specimens showing the fore
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19661 Carpenter - Protorthoptera and Orthoptera 79 Text-figure 16.
Paroedischia recta, n. sp. Drawing based on the holo- type (middle part of wing), paratype No. 15638 (base of wing) and para- type no. 15757 (apex of wing).
wings and legs. As noted above, the hind wings are unknown in the family7; a small and apparently distorted fragment of a hind wing is preserved in the type of Permoedischia moravica Kukalovi, from Czechoslovakia, but this genus can only doubtfully be referred to the Oedischiidae. It has been assumed, probably correctly, that the oedis- child hind wings had an expanded and folded anal area. I have dis- cussed above the peculiarities of the topography of the venation of Oedischia and have given reasons for my present belief that in this family, at least, the short vein connecting the stem of M to CuA is a modified cross vein, not a branch of M or a part of CLIA. Genus Paroedischia, new genus
Fore wing: shape much as in Metoedischia. Sc extending well beyond mid-wing, longer than in Metoedischia; Rs arising at about mid-wing, free piece of Rs (before anastomosis with M) longer than in Afeloedischia; separation of CuA and CUP much later than in Meloedischia; cross veins as in Metoedischia, but without formation of reticulation. Hind wing unknown.
Type-species : Paroedischia recta, n. sp. On the basis of the fore wing, which is all that is known, this genus seems to be close to Metoedischia but differs in the several respects already noted.
Paroedischia recta, n. sp.
Text-figure 16 and Plate 7
Fore wing: Length of holotype fragment, 20 mm.; width 7 mm.; estimated complete length of wing, 37 mm.; width 7 mm. Precostal space with a series of nearly parallel veinlets, connected by short 'The restoration of the oedischiid genus Metoedi~chia given by Martynov (1938, p. 49) was based on specimens of two species, one of which is now placed in the genus Pinegia of the family Tcholmanvissiidae.
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80 Psyche [March
cross veins; costal area with a series of straight or nearly straight veinlets, only very rarely branched; RI with several branches distally; Rs with several main branches, each forked; M dividing well before origin of Rs; MP sigmoidally curved; CuA with 6 terminal branches; space between IA and 2.A widened distally; cross veins as shown in figure .
Holotype: No. 58g7ab, M.C.Z., collected by F. M. Carpenter, in lower insect layer at Elmo, Kansas; this consists of the middle part of a well preserved fore wing. Several additional specimens almost certainly belonging to this species are as follows: No. 15757, Pea- body Museum, Yale University (collected by C. 0. Dunbar), con- sisting of a distal third of a fore wing; No. 15638ab, Peabody Museum (collected by C. 0. Dunbar) ; consisting of the basal quar- ter of a fore wing; No. 5900, M.C.Z.
(collected by F. M. Car-
penter), distal half of fore wing; No. 5898, M.C.Z. (collected by F. M. Carpenter), basal third of fore wing; No. 5899, M.C.Z. (col- lected by F. M. Carpenter), distal fragment of fore wing; No. 5896, M.C.Z. (collected by F. M. Carpenter), distal fragment of fore wing.
The fragmentary nature of these specimens is strongly indicative of unusually delicate wings, almost certainly membraneous, rather than coriaceous. A composite drawing of the fore wing of recta is included in figure1 16; the central part of the wing is drawn from the holotype; the basal and distal portions are based on specimens numbered I 5638 and I 5757, respectively. The venation of this insect is clearly subject to much variation, the number and precise arrangement of branches being different to some degree in all specimens.
Text-figure 17. Paroedischia maculata, n. sp, Drawing of fore wing (holotype) .
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Psyche
[March
Paroedischia maculata, n. sp.
Text-figure I 7
Fore wing: length, as preserved, 20 mm. width, 7 mm.; estimated complete length, 30 mm. Venation so far as known like that of iccta, except that the main veins are almost consistently forked at the wing margin, the branching of R is less, the main veins are thicker, and the cross veins are thinner and more irregular. In addition the wing has several distinct maculations, as shown in the photograph, plate 4. The wing of recta shows no signs of markings of any kind. Holotype: No. 5873ab, Museum of Comparative Zoology (Col- leered by F. M. Carpenter), in the lower layer of the Elmo lime- stone; this consists of a distal two-thirds of a wing, which is somewhat wringled along the posterior border but otherwise very well preserved. Family Permelcanidae Sharov
Permelcanidae Sharov, 1962, p. 112.
Fore wing: membranous or very weakly coriaceous. Precostal area well developed but not forming a prominent bulge, extending about one-fourth wing length from base; Sc extending slightly be- yond mid-wing; R distinct from Sc basally; Rs arising near or just beyond mid-wing, usually anastomosing with a branch of MA for at least a short distance; Rs and M with at least two terminal branches ; CLIA as in the Oedischiidae, with several terminal branches; CUP close to IA; IA with several veinlets leading to the hind mar- gin; 2A and 3A much shorter. Cross veins much fewer than in the Oedischiidae, not forming a true reticulation. Specimens showing both fore and hind wings have not been found, but isolated portions of hind wings, consisting mainly of the remi- gium in each case and probably belonging to this family, have been described by Sharov. In these hind wings the costa is short and submarginal; Sc, RI, Rs and M are essentially as in the fore wing; an anal area was piesumably present although only a suggestion of one is visible in the fossils. The body structure is unknown. Three previously described genera belong here: Promartynovia Tillyard, from the Elmo limestone, Kansas; and Permelcam Sharov and ProcZcma Sharov, from upper and lower Permian, deposits, re- spectively, in the USSR.
Genus Promarty novia Tillyard
Promaftynovia Tillyard, 1937, Amer. Journ. Sci., 33 :99. Fermelcana Sharov, 1962, Paleont. Journ. 2 :114. Fore wing: costa usually consisting of two or more distinct
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19661 Carpenter - Protorthoptera and Orthoptera 83 branches arising from the wing base; Rs arising just before the end of Sc; CuA with several terminal branches; IA extending well beyond 2A and sending several short branches to the hind margin; cross veins usually straight, only rarely branched. Pterostigmal and apical areas of wing pigmented.
Type-species :
Promarty n ovia venicosta Tillyard.
I have been unable to find sufficient differences between Pro- martynovia Tillyard and Permelcana Sharov to justify generic sep- aration. Prornanynovia was very incompletely known to Tillyard, who placed it in the order Neuroptera, and his description of P. venicosta was unsatisfactory, for the reasons given below. The type- species (sojane-rise Sharov) of Permelcana Sharov is known by a nearly complete fore wing, which lacks only the apical region. Sharov had little reason to associate his fossil from the Upper Permian of the USSR with Promartynovia, although I had pointed out in 1943 (p. 61) that the latter genus was orthopteroid not neuropteroid. I strongly suspect, also, that Proelcana Sharov, based on an apical frag- ment ( uralica Sharov ) from Lower Permian deposits of Tchekarda, USSR, will turn out to be a synonym of Proinartynov'ia; the amount of anastomosis between Rs and M, and the detailed arrangement of the branches of these veins, used by Sharov as generic characters, are highly variable within this group of orthopteroids. Promartynovia venicosta Tillyard
Text-figure I 8
Promarfynovia venicosia Tillyard, 1937, Amer. Journ. Sci. 33 :100; fig. 6 (Order Neuroptera, family Martynoviidae). Fore wing: length I I mm.; width 2.6 mm.; front margin ap- parently slightly concave; costa consisting of two main branches leading from wing base, the posterior one simple, the other forming Text-figure 18. Promartynovia vcnicosta Tillyard. Drawing of fore wing, based on holotype in Peabody Museum, Yale University.
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84 Psyche [March
a complex of several branches; costal veinlets oblique, long, un- branched; Rs with two branches; MA anastomosed with Rs for a short distance ; MP apparently unbranched. Holotype: No. 15594, Peabody Museum, Yale University. This specimen, when studied by Tillyard, had not been completely cleaned; it was known to him only by the base of the wing. Removal of mat- rix has revealed the greater part of the rest of the wing, as shown in figure 18; some parts of the posterior margin of the wing are missing but they are not critical for the determination of the species. In his figure of the fossil, Tillyard omitted the weakly developed CUP, which arises from Cu basally and runs closely parallel to IA. It can be seen clearly in the specimen, however, with the aid of low- angle illumination. Tillyard's assignment of Promartynovia to the Neuroptera (Sialoidea) was made with some doubt; actually he placed the genus in the family Martynoviidae, now included in the extinct order Diaphanopterodea. The similarity of the fore wing of venicosta to that of sojanense Shai-ov is really striking. They are
of comparable size, have a similar venation and even possess the identical pigmentation of the pterostigmal area. Family Parelcanidae, new family
Fore wing : more coriaceous than in Oedischiidae; precostal area forming a prominant bulge; Sc apparently extending well beyond mid-wing; Rs (so far as known) arising slightly beyond mid-wing, anastomosed with a branch of
M for a short distance; CuA and
CUP as in Oedischiidae;
IA nearly straight; cross veins about as numerous as in Oedischiidae. Hind wing and body unknown. This family, which is apparently more closely related to the Oedischiidae than to the Permelcanidae, is characterized mainly by the prominance of the precostal area and by the coriaceous nature of the fore wing.
Genus Parelcana, new genus
Fore wing: costa usually consisting of two distinct branches, the anterior one giving rise to a series of radiating veins, the posterior one forked; at the level of end of precostal area, the costal space is about as wide as the space between R and Sc; termination of IA close to end of 2A; cross veins close together, uniformly distributed; no reticulation formed (at least in basal half of wing). Type-species : Parelcana dilatata, n. sp.
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Carpenter - Protorthoptera and Orthoptera IA CUP CU A
Text-figure 19.
Parelcana dilafata, n. sp. Drawing of fore wing (holo- type).
Parelcana dilatata, n. sp.
Text-figure 19
Fore wing: length of preserved part 19 mm.; width, 5 mm.; estimated length of complete wing:, 36 mm.; costal veinlets slightly oblique, mostly unbranched; branches of CuA short. Details of vena- tion shown in figure 19.
Holotype: No. 6304 ab, Museum of Comparative Zoology (col- lected by F. M. Carpenter in lower layer of Elmo limestone). This type consists of a well preserved basal half of a fore wing. The precostal area of this wing is unusually prominent. Although the wing is incomplete, that part which is preserved seems to be definitely coriaceous; the cross veins are preserved as distinct ridges, which are especially prominent when observed under oblique light. Genus Petrelcana, new genus
Fore wing: distinctly coriaceous; slender. Precostal area not so long or so broad as in Parelcana, but projecting beyond the line of the anterior margin of the wing; costa little-known, irregularly formed; RI with several oblique, almost longitudinal veinlets be- yond the end of Sc; at level of end of precostal area, costal space much wider than space between Sc and R; termination of IA well beyond the end of 2A; cross veins more widely separated than in Parelcm and more irregular, forming a coarse reticulation in parts of the anal area.
Type-species : Petrelcana elongata, n. sp. Petrelcana elongata, n. sp.
Text-figure 20
Fore wing; length, as preserved, 33 mm.; width 7 mm.; estimated complete length, 40 mm.; veinlets widely separated and irregular in
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Psyche [March
Text-figure 20.
Petrelccna clongata, n. sp. Drawing of fore wing (holo- type).
most of costal area; cross veins between RI and Rs sigmoidal and parallel; branches of CuA long, extending beyond the level of anas- tomosis of Rs with M.
Holotype : No. 6306, Museum of Comparative Zoology, collected by F. ^\I. Carpenter in the lower layer of the Elmo limestone. This consists of a nearly complete fore wing, lacking only the apical region. The fossil is very well preserved and shows definite indications of pigmentation along almost the entire length of R and RI, the pig- mentation broadening: out to include surrounding areas after the origin of Rs. The precostal area is not entirely known, although in the specimen it clearly protects beyond the rest of the wing mar- gin; whether or not it forms as noticeable a bulge as in Parelcana cannot be determined. The family assignment of this genus may need to be changed when the form of the costa is known. BOLTON, H.
1925. Insects
Insects,
CARPENTER, F. M.
I
from Coal Measures of Comrnentry. Brit. Mus., Fossil 2 :1-56.
The Lower Permian insects of Kansas. Part 7. The Order Protoperlaria. Proc. Amer. Acad. Arts Sci. 70 :103-146. The Lower Permian insects of Kansas. Part 9. The Orders Neuroptera, Raphidiodea, Caloneurodea and Protorthoptera, with additional Protodonata and Megasecoptera. Proc. Amer. Acad. Arts Sci. 75 :55-84.
The Lower Permian insects of Kansas. Part 10. The Order Protorthoptera: the Family Liomopteridae and its relatives. Proc. Amer. Acad. Arts Sci. 78 (4) :185-219.
Key to extinct families of insects. In Brues, C. T., A. L. Melander and F. M. Carpenter, Classification of Insects. Bull. Mus. Comp. Zool. 108 :777-827.
COMSTOCK, J. H.
1918. The Wings of Insects. Cornstock Publ. Co.
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1898-99. The Wings of Insects. Amer. Nat. vols. 32, 33. (A series of 23 papers).
FRASER, F. C.
1931. A note on the fallaciousness of the theory of pretracheation of the wing venation of Odonata. Proc. Roy. Ent. Soc. London, 13: 60-70.
GUTHOKL, P.
1936. Neue Beitiage zur Insekten-Fauna des Saarcarbons. Sencken. 18 : ZIb. 72 :1-51.
HAGEN, H. A.
1870 Ueber rationelle Benennung des Geaders in den Fliigeln der Insekten.
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HAUPT, H.
1941. Die altesten geflugelten Insekten und ihre Beziehungen zur Fauna der Jetztzeit. Z. Naturw. 94: 60-121.
HENKE, K.
1953. Die Musterbildung der Versorgungssystem in Insektenfliigel. Biol. Zlb. 72 :1-51.
HOLDSWORTH, R.
1940. Histology of the wing pads of the early instars of Pteronarcys froteus Newman. Psyche, 47 :112-120.
1941. The wing development of Pteronarcys proteus Newman. Journ. Morphology, 70 :431-461.
KUKALOVA', J.
1959. On the family Blattinopsidae Bolton, 1925. Roz. ceskosl. acad. vid., 69 (1) : 1-27.
1965. Permian Protelytroptera, Coleoptera and Protorthoptera (Insects) of Moravia. Sbornik geol. vid. pal. 6:61-98. LAMEERE, A.
1922. Sur la nervation alaire des insectes. Bull. Class Sci. Belgium, 1922 :138-149 (transi. Psyche 30 :123-132, 1930). LESTON, D.
1962. Tracheal capture in ontogenetic and phylogenetic phases of in- sect wing development. Proc. R. Ent. Soc. Lond. (A) 37:135-144. MARTYNOV, A. V.
1924. Sur ]'interpretation de la nervuration et de la tracheation des ailes des Odonates et des Agnathes. Rev. Russe d'Ent., 18:145-174. (transl. Psyche 3 7 :245-280, 1930).
1937. Permian fossil insects from Kargala and their relationships. Trav. Inst. Paliont. Acad. Sci. URSS, 7:l-92.
1938. Etudes sur i'hiiitoire giologique et de phyloginie des ordres des insectes (Pterygota). Trav. Inst. Paliont. Acad. Sci. URSS 7: 1-150.
NEEDHAM, J. G.
1935
Some basic principles of insect wing-venation. Journ. N. Y. Ent. SOC. 43 :113-129.
1951. Prodrome for a manual of the dragonflies of North America, with extended comments on wing venation systems. Trans. Amer. Ent. å£oc 77:21-62.
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88 Psyche [March
REDTENBACHER, J.
1886. Vergleichende Studien iiber das Fliigelgeader der Insekten. Ann. des k.k. nat. Hofmuseums 1. (3) : 153-232. ROHDENDORF, B. B., ET AL.
1962. Osnovy Paleontologii. Insecta. pp. 1-373. Akad. Nauk. SSSR. SCUDDER, S. H.
1885. Palaeodictyoptera: or the affinities and classification of Paleozoic Hexapoda. Mem. Bost. Soc. Nat. Hist. 3 :319-351. SELLARDS, E. H.
1909. Types of Permian insects. Part 3, Amer. Journ. Sci. 27:151-173. SHAROV, A. G.
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On the system of the Orthopterous Insects. Internat. Cong. Ent., Wien, 1960 (1) : 295-296.
1962. A new Permian family Permelcanidae (Orthoptera). Pal. Journ. 2 :112-116.
SMART, J.
1956. A note on insect wing veins and their tracheae. Quart. Jour. Micros. Sci. 97 (4) : 535-539.
SPEITH, HERMANN.
1932. A new method of studying the wing veins of the mayflies and some results obtained therefrom.
Ent. News, 43:103.
TILLYARD, R. J.
1923. The wing-venation of the order Plectoptera or mayflies. Journ. Linn. Soc. Zoology, 3 5 :143-162.
1927. Kansas Permian insects. Part 17. The Order Megasecoptera and additions to the Palaeodictyoptera, Odonata, Protoperlaria, Copeognatha and Neuroptera. Amer. Journ. Sci. 33 532-110. 1928. The new order Protoperlaria. Amer. Journ. Sci. 16 :l85-220. WATERLOT, G.
1934. Bassin houiller de la Sarre et de la Lorraine. ii. Faune fossile. Etudes Gites Min. Fr: 1934. 111-221; 269-273.
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