Jarmila Kukalová-Peck.
Unusual Structures in the Paleozoic Insect Orders Megasecoptera and Palaeodictyoptera, with a Description of a New Family.
Psyche 79(3):243-268, 1972.

This article at Hindawi Publishing: https://doi.org/10.1155/1972/98019
CEC's scan of this article: http://psyche.entclub.org/pdf/79/79-243.pdf, 3936K
This landing page: http://psyche.entclub.org/79/79-243.html

The following unprocessed text is extracted automatically from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

UNUSUAL STRUCTURES IN
THE PALEOZOIC INSECT ORDERS
MEGASECOPTERA AND PALAEODICTYOPTERA,
WITH A DESCRIPTION OF A NEW FAMILY*
BY JARMILA KUKALOVA'-PECK
Department of Geology, Carleton University, Ottawa, Ontario, Canada
The order Megasecoptera is a representative of the haustellate paleopterous insects of the evolutionary line that lived during the Pennsylvanian and the Permian. The similarity of wings and body structures, such as mouth-parts and genitalia, indicate very close re- lationship with the order Palaeodicty~~tera. Both groups are pre- sumed to have emerged sometime during the Mississippian from a common ancestor. While Palaeodictyoptera are usually larger and more sturdily built, bearing broad wings with a rich venation and prothoracic lobes, Megasecoptera are slender insects with a more delicate appearance, with petiolate wings and simplified venation, with enlarged thorax lacking prothoracic lobes, and with tapering abdomen.
The present paper deals with an extraordinary morphological feature - projections of the body cuticle, which occur in most or all Megasecoptera and at least in some Palaeodictyoptera. These are conspicuous processes, which are short to very long, simple or branched, and which are distributed in regular rows on the ab- domen and thorax.
A fuller understanding of the morphology of this very unusual character resulted from two years of intensive research by Dr. 3'. M. Carpenter and myself, based upon fossil material of Commentry (Upper Pennsylvanian, France), Mazon Creek (Middle Pennsyl- vanian, Illinois), Obora (Lower Permian, Czechoslovakia), Elmo (Lower Permian, Kansas), and now also Tshekarda (Lower Per- mian, Siberia). I am deeply indebted to Professor Carpenter, who was very helpful in the preparation of this study. Until now, the projections have been only poorly known. They were at first mostly interpreted as tracheal gills that persisted into the *This study has been supported in part by grant No. GB-27333 (F. M. Carpenter, Principal investigator, Harvard University) from the National Science Foundation.
Manuscript received by the editor July 15,1972. Pnrfif 79:MJ-268 Mil}. http://psyche cnlclub w@9#Ìö html



================================================================================

244 Psyche
[September
adult stage, or later as short lateral spines on the abdominal segments and thorax. The following is a short account of present knowledge. The projections on the ~rothorax are, in some families, long and filiform, but in other families rather short, pointed and spine-like. They were described as spines in some Megasecoptera, namely in Mischoptera, Aspidothorax, Corydaloides and Foriria by Brongniart ( 1885 ab, 1890, 1893), Lameere ( 1908, 191 7), Carpenter ( 1951, 1968) and others. In 1968 Carpenter and Richardson mentioned stout lateral spines in the nymph of Mischoptera douglassi on the meso- and metathorax.
The abdominal projections are actually filiforrn, growing in fringe- like rows out of the tergites. However, all previous authors ob- served only the basal parts of several abdominal projections situated laterally, which led to incorrect interpretations. Thus Brongniart ( 1885, p. 63 ; 1885, p. 658; 1890, p. I 540) considered them to be branchio-tracheal appendages, which served for aquatic respiration in nymphs and which were carried over to the adults. In his general account on Carboniferous insects of Commentry, he gave a detailed figure ( 1893, p. 305, p. 298, fig. 50) of an enlarged "lateral lamella" with branched "tracheae" in the genus Corydaloides (Mischopte- ridae). His point of view was followed by Brauer (1886, p. 107), who classified the projections as "persistent abdominal tracheal gills". Handlirsch first ( I 906) stated that Megasecoptera possessed "den- tated lamellar appendages, which were perhaps derived from tracheal gills".
The gill character of the projections was denied by Lameere ( 1908, p. 136; 1917, p. 28; 1917, p. 145), who compared the "lamellae" with the lateral expansions of the Recent mayfly Onisciguster wake- fieldi (N. Zealand). He regarded the projections protruding out from "lamellae" to be backwardly directed spines. Martynov (1938, p. 25) characterized Megasecoptera as having "lateral expansions of abdominal segments with tooth-like or spine- like outgrowths, homologous with prothoracic spines and prothoracic winglets of Palaeodictyoptera, reduced and modified". Carpenter (195 I, p. 353) correctly stated that the projections were extensions of tergites, but also believed them to be short and spine-like in char- acter (Corydaloididae, 195 I, p. 35 I ). A significant step in the research of the character of abdominal projections was the paper on megasecopterous nymphs published by Carpenter and Richardson ( I 968 ) .
In this remarkably preserved
nymph, Mischaptera douglassi, the hind margins of the abdominal tergites, except the last two, bear a row of seven stout "spines". This



================================================================================

19721 Kukalovii-Peck - Palaeozoic Insect Orders 245 fact was unusual enough to inspire the thoroughfull examination of abdominal tergites of all known Megasecoptera for this feature. After a detailed discussion with Dr. Carpenter (during my tenure as Alexander Agassiz Lecturer in Zoology at Harvard University), I visited the Museum d'Histoire Naturelle in Paris on my return trip to Europe. This institution's collections hold the most extensive material of Paleozoic Megasecoptera (Brongniart, 1893 ; Carpenter, I 95 I ) . I found that each of the sufficiently preserved megasecopteran bodies (mostly Mischopteridae) had prolonged filaments leading from the posterior margin of the abdominal tergites. The projections were visible only under glycerin, a medium which was o~bviously not ap- plied to the fossils by previous students. It should be noted that in the Mischoptera douglassi nymph the bases of the projections give a perfect spine-like appearance, which now seems to be due to incom- plete preservation. Recently, Carpenter and Richardson ( 197 I ) de- scribed long filamentous projections in Eubrodia dabasinskuxi (Brodi- idae) extending posteriorly along the mesothorax to almost the end of the body.
The specimens of Megasecoptera and Palaeodictyoptera newly in- troduced in the present paper contribute significant features to the knowledge of the projections. Sylvohymen sibiricus nsp. (Bardohy- menidae), a megasecopteron from the Lower Permian of Siberia, shows the hollow, broken bases of projections located not only along the posterior margin of abdominal tergites, but also on tergal nota of the whole body (fig. I and pi. I ). M~onsteropterum moravicum nsp., a palaeodictyopteron from the Lower Permian of Czechoslo- vakia, presents well preserved projections (fig. 6 and pi. 3)) showing details of the surface and of multiple branching. Summarizing our present knowledge, we can say that the processes or projections are hollow outgrowths of the tergites and are usually arranged into regular transverse rows, are simple or branched, and are short to very long, according to the particular families. The outgrowths are directed up and backwards from the body, so that they protrude. The ventral side of the ~rojection-bearing bodies is not known. On the thorax, the ~rojections may form spines, or may be filiform, identical to those on the abdomen. The abdominal ~rojec- tions with their superimposed series of fringes, resemble the tradi- tional skirt of the Spanish national female costume. It is possible that all species of Megasecoptera possessed projections, more or less developed, both adults and nymphs. Projections of the same char- acter occurred in some Palaeodictyoptera, but probably not in all families.




================================================================================

246 Psyche [September
It has taken almost 80 years for acquisition of the above data about these two groups of insects, though they are not rare in Paleozoic deposits. This slow process becomes more understandable after con- sidering the character and nature of preservation of the projections. Protruding above the body in life, they tended to become hidden by the superimposed sediment rather than become compressed at the same level as the body itself during fossilization. The broken and usually more sclerotized bases of the projections are not distinguishable from spines or tubercles and are mostly inconspicuous. Finally, in a ma- trix which does not preserve chitin, the imprints of the projections are vague.
During my study, the projections were thoroughly examined for connections with the insect's body. In the matrix capable of pre- serving chitin (e.g. Cornmentry shale), the projections have the same brown color as the terga. Their surface is covered with a rugosity similar to that on the body (Mazon Creek, Illinois) or with a minor rugosity and scattered sockets of setae (Obora, Moravia). In Mon- steropteruin moravicum, the surface of the projections is identical to that of the legs. The arrangement of the projections is regular and probably characteristic for all genera within a family (I have found this to be true so far for Bardohyrnenidae, Protohymenidae and Mis- chopteridae). The width of the projections varies with the size of specimens; their arrangement is bilaterally symmetrical. The above mentioned features exclude the possibility that the outgrowths are parasitic organisms or fungi.
These projections in the Megasecoptera and Palaeodictyoptera ap- pear unique among insect orders, and their function remains ob- scure. However, several features suggest that they might be homo- logous to certain tergal structures of Odonata. In all Recent Odo- nata, there is a transverse ridge at each end of the tergum, the anterior and posterior transverse carinae (Walker 1953, p. 18). The former is inconspicuous, but the posterior carina is a distinct ridge bearing a row of small tubercles or denticles. By the position and arrangement in rows, the projections in Megasecoptera and Palaeodic- tyoptera are very suggestive of the prolonged and enlarged carinal denticles of Odonata. Their function, of course, presents a com-
plicated problem, which can hardly be solved wi'th fossil material. ORDER MEGASECOPTERA
Family Bardohymenidae Zalessky
Type Genus: Bardohymen Zalessky, 1937.
This family was erected by G. Zalessky ( 1937) and redefined by



================================================================================

Figure 1. Syloohymen s'tbiricu~ n.sp.; Pr = bases of hollow outgrowth of termites. Holotype: fore wing length 50 mm, width 9.1 mm. Lower Permian of Siberia.



================================================================================

248 Psyche [September
Carpenter (1947). A significant contribution to the morphology of the wings was published by Carpenter (1962). Until now, the family has been based only upon isolated, more or less fragmentary wings.
The following introduces certain characteristics of body structures and some additional features of wing morphology based upon 5 new specimens from the Lower Permian of Siberia and Czechoslovakia. Wings subequal in length and shape, strongly petiolate, similar in venation; C flattened and wide, very close to Sc; Sc distinguishable as a separate vein only in proximal part of the wing; RI contiguous with C and Sc except in the very distal part of the wing; RI with short terminal branches; Rs originating at about midwing, giving rise to 2-3 branches; M very close to R basally, diverging away from R beyond the first quarter of the wing length; M dividing into MA and MP at variable level, but near to the origin of Rs; MA con- nected with Rs or R with a strong cross vein; 'Cu at the base fused with the stem of M; CuA connected with the stem of M by a strong cross vein; 2 anal veins, A1 long with a pectinate series of branches; A2 very short and simple; cross veins not numerous, usually ar- ranged in 2 rows; veins and wing margin with rows of setal bases or sockets.
Body structures: head small, short and broad, with large pro- jecting eyes ; antennae long, composed of many cylindrical segments ; maxillary palpi robust; prothorax trapezoidal; mesothorax and meta- thorax large in proportion to the rest of the body; legs of middle length, cursorial; abdomen relatively slender tapering abruptly in the anterior part; females with I I visible segments and protruding ovipositor; projections forming rows on the posterior margin of thoracic and abdominal segments; parallel, transverse rows of projec- tions on abdominal terga, and occasionally on thoracic segments ; larger projections located in pairs in the central parts of the body segments. The family Bardohymenidae is closely related by wing morphology to Protohymenidae (Carpenter, 1962)~ which turns out to be true also for the body. However, the wing venation is less advanced, possessing an MA which is not anastornosed with Rs, and a CuA free from M. Also the general form of the wings is less specialized, as the hind wings are almost equal to the fore wings, not reduced in length as in the Protohymenidae. The body in both families is much alike, possessing a large thorax and tapered abdomen. The bardohy- menid body is, in relation to the wings, more heavy. Through the courtesy of Dr. Carpenter I was able to study Protohymen readi Carpenter (1933), Protohymen elongatus Carpenter (1930) and



================================================================================

19721 Kukalovd-Peck - Palaeozoic Insect Orders 249 Prwtohymen j%m-mianus Tillyard ( I 924) for projections. In all three specimens the bases of projections are present and very similarily dis- tributed as in Bardohymenidae. However, they merge with the un- even surface of
the rock to such an extent that they would be undetectable unless a well preserved specimen, such as the type of S. sibiricus n.sp., were available for comparison. By delicate prepara- tion of the surrounding matrix I was able to1 uncover remnants of projections (P. readi, specimen 3257, Museum of Comp. Zoology, Harvard University; P. permianus, specimen 5053, Peabody Museum, Yale University), which are prolonged and backwardly curved. This fact is very significant, because in Sylvohym-en sibirici~s nsp. (Bardo- hymenidae) the projections continue into the covering matrix and cannot be followed.
The projections in Bardohymenidae and Protohymenidae are ar- ranged in transverse rows. By position and distribution they are very similar to denticles in the transverse carinae of Odonata. In my opinion, these structures may be homologous. Besides, some anisop- teran nymphs (for instance Erpetogomphus designatus, Gomphidae) Needham & Westfall, 1955, bear, on several abdominal terga, paired darker pits, located along the median line precisely like the bases of the large paired projections in Bardohymenidae and Protohymenidae. This similarity is suggestive of ~ossible musculature inside the paired projections in Megasecoptera.
Genera included: Bardohymen G. Zalessky, 1937 (Lower Per- mian, Barda River, U.S.S.R.) ; Sylvohymen Martynov, 1941 (Lower Permian, Tshekarda, Siberia, U.S.S.R. and Lower Permian, Okla- homa) ; Calohynzen Carpenter, 1947 (Lower Permian, Oklahoma) ; Actinohymen Carpenter, 1962 (Lower Permian, Texas) ; Alexahy- men n.g. ( Lower Permian, Czechoslovakia). Genus Sylvohymen Martynov
Sylvohymen Martynov, 1941: 10; Carpenter, 1947: 31; Carpenter, 1962: 37; Rohdendorf, 1962: 68.
Type species: Sylvohymen robustus Martynov, 1938 (OD). This genus is based upon a distal part of a wing from the same locality (Tshekarda) as the presently described specimen of Sibiricus, n.sp., which is much more fully preserved. Carpenter ( 1946, p. 31, fig. 7) described 8. ingens from the Lower Permian of Oklahoma, also based on a distal wing third. The apical parts of the wing in all three specimens resemble each other and the species cannot be separated generically as far as is presently known. Unfortunately, they can also hardly be separated from Bardohymen (this statement



================================================================================

Psyche [September
PLATE 1
SyIwohyflies. sibwicus n,ap,, obverse. Vie arrows point to the banes of the proJcctioDs on the tergite~. Lower Permian, Siberia. is based MI literature only) and further study of the original mate- rial might find the two genera synonymous. Martyonv's reconstructib of Sylvohymen robustsus ( 194 I ) show- ing MA anastornosing with Rs is obviously incorrect, as the connet- tion of MA with R or Rs by means of a cross vein is characteristic for Bardohymenidae.
Wings long and slender, tapered rather abruptly in the basal third; hind wings slightly longer than fore wings, broader at about mid- wing; Sc recognizable only in proximal half of the wing; Rs with 3 main branches; AI S-shaped; posterior margin in untapered part of wing almost parallel to the anterior margin. Body structures : Prothorax with transverse elevations ; rnetathurax broader than mesothorax; first abdominal segment strongly tapering;



================================================================================

19721 Kukalovd-Peck - Palaeozoic Insect Orders 25 1 ovipositor in females stout, covered by hairs; projections probably maximally 14 in number in a row.
As shown previously by Martynov ( 1941) and Carpenter ( 1947, p. 32)) Sylvohymen is closely related to the type genus Bardohymen. The reconstruction of Bardohymen magnipennifer (G. Zalessky 1937, p. 603, fig. I) is obviously incorrect for its cubital branches. By the structure of pterostigmal area the genus Sylvohymen is related to Alexahymen n.g., which differs in having a relatively shorter and broader wing with concave posterior margin, small rs a;-ea and AI parallel with posterior margin.
Species included : Sylvohymen robustus Martynov, I 938 (Lower Permian, Oklahoma) ; Sylvohymen sibiricus n.sp. (Lower Permian, Tshekarda, Siberia).
Sylvohymen sibiricus n.sp.l
Figure I, plates I and 2
This species is based upon an obverse and reverse of a female with two complete wings and damaged lateral part of body. The thorax and the abdomen are preserved on the dorsal side, while the head pre- sents a composite of dorsal and ventral surface showing bases of stout palpi. The projections, if only the obverse were known, give the appearance of stout tubercles. In the reverse, however, they con- tinue like hollow tubes into the matrix. For preparing the illustration, both obverse and reverse parts of the specimen were used.
Wings slightly subequal, the hind pair being longer and broader at about midwing; color markings missing; fore wing length 50 mm, maximum width 9. I mm, almost equally broad except for the tapered proximal third; anterior margin slightly convex; C bordering the whole wing; apex bent backward and almost pointed; RI apically diverging to some extent, with 1-3 terminal twigs; Rs with 3 simple long branches; A1 S-shaped with a row of about 9 branches; cross veins about 18 in number; cross vein between RI and Rs forming a heavy bar, thickened at its costal end; hind wing length 51 mm, maximum width 9.7 mm broadest at the mid-wing; hind wing nar- rowing proximally less abruptly; anterior margin somewhat straighter ; RI apically less diverged away from the anterior margin, with only I twig.
'This remarkable specimen was turned over to me for study by the courtesy of Dr. B. B. Rohdendorf, the head of the Paleoentomological De- partment of the Paleontological Institute of the Academy of Sciences in Moscow, for which I express my sincere gratitude.



================================================================================

Psyche
[September
PLATE 2
Syiwokymen ~ihiricus n.sp., reverse. The veins are secondarily colored by manganese. Arrows point to hollow continuations of projections into the matrix. Lower Permian, Siberia.




================================================================================

19721 Kukalovd-Peck - Palaeozoic Insect Orders 253 Body structures: Length of head 1.8 mm, width about 3.6 mm; preserved length of antennae 9 mm, antennae composed of numerous cylindrical segments ; segments of maxillary palpi striated, p reserved segment length 1.1 mm, width 0.8 mm; median line running along the whole body; prothorax length 4 mm, maximum width 8 mm, provided with two obliquely oriented longitudinal elevations and one elevation located anteriorly and centrally; mesothorax length 4.1 mm, probable width 8.8 mm; metathorax length 4.2 mm, maximum width probably 9.2 mm; legs covered by setae; front tibia length 38 mm; hind tibia length about 50 mm; abdomen length 14.5 mm, maximum width 9.4 mm; abdominal segments unequal, length of segments as follows : 1st 2 mm; 2nd 0.8 mm ; 3rd I .3 mm ; 4th I .7 mm ; 5th 2 mm; 6th 2 mm; 7th 0.5 mm; 8th I mm; 9th I mm; 10th 1.6 mm; I ith 0.5 mm; each abdominal segment but the I ith has a transverse flat topped ridge; I ith segment divided by a deep incision into' two lobes; ovipositor stout, reaching much beyond the end of the body, covered by dense stiff hairs oriented anteriorly. Projections: Two stout projections located in the central part of each body segment except the I I th; prothorax with an additional pair of projections anteriorly and with about 8 projections along the pos- terior margin; mesothorax with a row of small projections parallel and near to the anterior margin and with double row of stouter projections on the posterior margin; metathorax with a series of stout projections on posterior margin; abdominal segments with a row of stouter projections on the flat topped ridge and another row of smaller projections bordering the posterior margin; I I th segment
with only the posterior row of minute projections. Holotype : No. I 700/394, Paleoentomological Department, Paleon- tological Institute of the Academy of Sciences in Moscow. Collected in Lower Permian deposits of Tshekarda, Siberia. The preservation of the holotype is very good, particularly be- cause the veins of the wings have been secondarily penetrated and colored by manganese, which enters also the minute transverse cracks. The body is not fully flattened. The abdomen especially is preserved in its original convexity. Some of the bases of the projections are well preserved and only those are introduced in figure I, marked as circles, as they actually appear. The projections were undoubtedly growing out from the tergites in regular rows, but since the surface of the body is uneven, they cannot be distinguished from the irregu- larities of the matrix. The actual length of the projections could not be followed as they continue inside the reverse of the fossil under an acute angle with the body. Their position, however, indicates that



================================================================================

254 Psyche
[September
they have been sclerotized. In analogy to the closely related Protohy- menidae, it is probable that the projections were slightly curved and at least several millimeters long.
Genus Alexahymen, new genus
Type species': Alexahymen maruska n.sp., Lower Permian of Moravia. This genus includes one species, represented by 3 incomplete wings. Wings shorter than in the related genus Sylvohymen, broadest be- hind the midwing, tapering gradually towards the base; Sc distinct to about two thirds of the wing length; RI diverging apically slightly from the anterior margin; Rs with two short branches; A1 parallel with the posterior margin; posterior margin slightly concave. Alexahymen differs from other genera of the family Bardohy- menidae in its relatively short and broad, gradually tapering wings, posterior margin concavely shaped, A1
parallel with the posterior
margin and sending off a series of numerous twigs, and small rs area. Species included: Alexahymen maruska n.sp. (Lower Permian, Obora, Moravia) .
Alexahymen maruska n.sp.
Figure 2, 3, 4
Derivatio nominis: In honor of Mrs. Maruska Alexovi, who generously gave support and encouragement to workers at the Obora locality for ten years.
This species is based upon the holotype, represented by a wing without the proximal part, and by two additional, isolated, equally damaged wings. With regard to the close similarity between the fore and hind wings in Bardohymenidae, the position of wings in the pair can be only inferred. However, from analogy with Sylvohymen sibiri- cus, the only bardohymenid with both wings in situ, it seems that the hind wings in this family tended to have a straighter anterior margin and more concave posterior margin. Consequently the hole type (fig. 2) and specimen 2/1972 (fig. 3) are represented probably by the hind wing, while specimen 3/1972 (fig. 4) is more likely the fore wing.
Color markings missing; total wing length about 33-36 mm, maxi- mum width 8.7-9.9 mm; anterior margin slightly convex in the distal part; C bordering the wing; apex more or less pointed; RI with 4-5 very short terminal twigs; Rs branches short; Ax sending off 6- branches; cross veins I 5-16 in number, mostly in double row. Material : Holotype No. I/I 972 (obverse and reverse probably of hind wing) ; specimen No. 2/1972 (obverse and reverse probably of



================================================================================

Kukalovd-Peck - Palaeozoic Insect Orders 255
Figure 2. Alexahymen maruska n.sp.; hind wing: length 30 mm, width 9 mm. Holotype. Lower Permian of Czechoslovakia. Figure 3. Alexahymen maruska n.sp.; hind wing: length 31.8 mm, width, 9.9 mm. Specimen 2/1972. Lower Permian of Czechoslovakia. Figure 4. Alexahymen maruska n.sp.; fore wing: len,gth 23 mm, width 8.7 mm. Specimen 3/1972. Lower Permian of Czechoslovakia.



================================================================================

256 Psyche
[September
hind wing) ; specimen No. 3/1g72 (obverse and reverse probably of fore wing) ; Paleontological Institute of Charles University, Prague, Czechoslovakia. Collected in the Lower Permian deposits near Obora, Moravia.
All three specimens of AZexahymen maruska carry the details of venation, described by Carpenter (1962, p. 38-39) in Actinohymen russeli, namely flattened C, widening distally beyond the end of Sc; and C, Sc, R-RI around midwing touching each other. The cross vein rs-r1 forms a heavy bar widened at its costal end in the holotype and specimen 3/1972. In the specimen 2/1g72 it is an average, though thick cross vein.
The posterior margin in the holotype is formed in a different way, perhaps as individual variation.
It is convexly curved in between the
branches of AI, media and cubitus, so that the tips of the branches protrude not unlike the fingertips in a bat wing. A similar phenome- non is indicated in Moravohymen vitreus n.sp. of the related family Moravohymenidae.
Moravohymenidae, new family
Type genus : Moravohymen n.g.
This family is based upon a fragment of a single wing (probably hind wing), which seems to combine the features of Bardohymenidae with those of some megasecopterid families of Commentry, France (Carpenter, 195 I ) .
Wings broadest at the beginning of the apical third, tapering grad- ually proximally; Sc remote from the anterior margin and terminating freely in the subcostal area well before apex; RI remote from Sc, not diverging apically from the posterior margin; MA connected with R or with the very origin of Rs by a cross vein; stem of M either close or fused with R; A1 not parallel with the posterior margin, sending off few irregular branches; cross veins arranged into irregu- lar rows and partly sigmoidal; row of cross veins in r1-rs area; veins and wing margin provided by setae.
The family Moravohymenidae has very gradually tapering wings with maximum width shifted to the distal third. MA is connected with R or Rs very much as in Bardohymenidae. The arrangement of the rest of the veins and sigmoidal cross veins reminds one more of some Upper Carboniferous Comrnentry families such as Mischop- teridae, Sphecopteridae and Corydaloididae. Genus included:
Moravohymen n.g. (Lower Permian, Czecho- slovakia).




================================================================================

Kukalov&Peck - Palaeozoic Insect Orders
257
Figure 5.
Moravohymen vitreus n.sp.; hind wing: length 21 mm, width 6.9 mm. Holotype. Lower Permian of Czechoslovakia. Genus Moravohymen new genus
Type species :
Moravohymen vitreus n.sp., Lower Permian of Moravia. Wings rather small, narrow in the proximal half, broad in the distal half ; C not flattened ; Sc rather a thin vein ; R i sending off several short twigs apically; Rs originating shortly behind midwing; Rs with 3 branches; branches of M and Cu simple; A1 remote from the posterior margin, anal branches irregular and rather long; cross veins arranged by two or three in the posterior part of the wing. Species included: Moravohymen vitreus n.sp. (Lower Permian of Obora, Moravia) .
Moravohymen vitreus nsp.
Figure 5
This species is based upon an obverse and reverse of a wing with damaged proximal part. According to the concave shape of the pos- terior margin it might be a hind wing.
Wing fragment: length 21 mm, maximum width 6.9 mm; C, Sc and RI distally much thinner veins than in Bardohymenidae; the membrane in the pterostigmal region probably sclerotized ; RI send- ing off 3 terminal twigs to C; SI-rs area broad, with 6 weak cross veins; Rs branches occupying a considerably large area; cross vein connecting MA with R at the origin of Rs is a heavy bar, thickened at its costal end; cross veins in medial area and cubital area slightly sigmoidal; posterior margin with small convex bends in between the ends of median and cubital branches.
Holotype: No. 4/1972 (obverse and reverse probably of hind wing) ; Paleontological Institute of Charles University, Prague,



================================================================================

Psyche [September
Figure 6.
Monstero$tcrum moravicum n.sp.;
C = coxa; L = lacinia;
0 = ovipositor; Pr == projections provided with setae; S = stylus; T trochanter. Original, ventral view. Lower Permian of Czechoslo- vakia.




================================================================================

19721 Kukdov&Peck - PaZmozoic Insect Order$ 25 9 Czechoslovakia. Collected in the Lower Permian deposits near Obora? Moravia.
ORDER PALAEODICTYOPTERA
Family Hom~io~teridae Handlirsch
Genus Monsteropterum, new genus
Type species:
Monsteropterum moravicum n.sp., Lower Permian of Mora- via.
This remarkable fossil with fragmentary wings would hardly war- rant formal description because the classification of the order is based upon the wing venation. However, the specimen shows the inner structure of the sucking mouth parts, the ventral attachment of the legs to the body, the branched ~rojections of terga with reserved surface and an ovipositor provided by styli. Since this insect is of unusual interest, generic and specific names are being assigned. Though the wings of the specimen are fragmentary, there is no doubt about referring them to the family Homoiopteridae, according to following characteristic features (I<ukalovi 1969, p. 440) : stem of main veins with a bend in the basal third of the wings; CuA and CUP parallel to each other; numerous, irregular and often connected cross veins.
Of the genera included, B~oZtopruvostia Strand, I929 is probably the nearest related genus. From this, Monsteroptemm dif- fers in lacking the sclerotized strip and tubercles, strengthening the costal area and in the more proximal division of M. The new genus is the first Permian representative of this rather primitive family and extends its occurrence from Upper Nmurian to Lower Permian. Monsteropterum moravicum n.sp.
Figures 6, 7, 8, 9, 10; plate 3
The body is a composite of ventral surface and inner structures, which were uncovered by preparation at different levels. With regard to the complexity of the compositej preservation must be discussed next.
The bod; was preserved while lying on its dorsal side. It was
much decomposed before being covered by sediment. The maxillary palpi disintegrated into single segments, which were partly displaced. The legs with some parts of the sterna were shifted towards the head. Valves of the ovipositor were split open, Only legs, wings, and one
segment of maxillary palpi show the natural ventral surface. The
beak split unevenly along the median plane, showing the inner side of two mandibular stylets and a small fragment of the distal end of one maxillary stylet (fig. 7-Ma). Meso- and metathorax expose the inner surface of the terga. The abdomen split between the sternal



================================================================================

Psyche
[September
PLATE 3
Monsieroptcrum momwicum n.sp.
Ventral side of the body showing long
beak, attachment of legs to the body and tergai projection9 (P). hwcr Permian, Cztchoslovakia,
and tergal elements and is vaguely pmerved. It was partly removed to uncover the projections.
Head structures:
The character of the pdaeodictyopteran mouth- - "
parts, which are eIongated into a beak co&sting of stylets of man- dibk and maxillae, have been dscribed in more detail by Crampton (1927) and by Laurentiaux ( 1952, 1953). However, there was no evidence about the arrangement of the stylets. Only recently Car- penter and Richardson ( 1971, p, 280) described a section of the beak in a strikingly unusuaIIy preserved specimen. The pair of some- Figure 7. Enlarged beak of Mon~tctopttrum tnorawirum n~p., ventral view. The beak split unevenly along median plane between two pairs of 3tyIeta. A43 = section of the beak figured on the bbk diagram in fig. 8; E = oval elevation; L = lacinia; Ma = ftagment of the maxilla^ stylet represented by the apical part of galea ; Md = natural inner surface of mandibular stykt where it contacts maxillary stylet; P z= crescent- ahaped pit. hwer Permian of Czechodovak!a.



================================================================================

Kukalovd-Peck - Palaeozoic Insect Orders 26 I




================================================================================

262 Psyche [September
what larger mandibles is located anteriorly from the maxillae and there is the fifth stylet, probably derived from the hypopharynx, be- tween the maxillae and slightly more posterior. The arrangement of the stylets in Monsteropterum mormicum fully confirms the conclusion of Carpenter and Richardson. The pair of mandibles is superimposed, and slightly longer than the pair of maxillae. The mandibular stylets in all probability partly overlap each other along the inner margin, because the undisturbed width of the left mandibular stylet (fig. 7-Md) extends much beyond the ideal median line. The same madibular stylet probably exposes in this part (fig. 7-Md) its natural inner surface where it contacts the maxillary stylets. It is provided by alternating ridges and grooves (fig. 8-R, G) probably enabling firmer connection in between stylets and strengthen- ing the long stylets.
The rest of the beak (more proximal and right part in fig. 7) shows the inside surface of the cavity in the mandibular stylets. It
seems certain that the mandibles were hollow, as in Recent dragon- flies (P. s. Corbet, in lift.). The mandibles of other extant insects are mostly solid except for occasional cavities and canals containing nerves for sensillae and haemolymph.
The ho1low nature of the
elongated mandibles of the Palaeodictyoptera may be explained as a means of reducing the mass of the head.
This assumption seems an
acceptable solution for the mechanical problems of flight engendered by the considerable weight of the head when compared to the rest of the body.
In the cavity inside the mandibular stylets, there are 5 rows of deep crescent-shaped pits (fig. 7-P; 8-P) and oval elevations (fig. 7-E; 8-E). They form continuous rows and seem to belong to a single structural unit, which is a series of short, peg-shaped, perpendicu- larly oriented pillars, supporting the long holloi mandibular stylets from inside. This assumption is based mainly on the fact that the second row of crescent-shaped pits (fig. 7-P) passes distinctly under the layer Md (fig. 7)) which is the natural surface of the mandible in contact with the maxillary stylet. On the Md layer the pit row continues in the form of oval elevations (fig. 7-E). The left maxillary stylet is completely missing; the right is prc- served only by the fragment of distal end (fig. 7-Ma). However, it provides information on the morphology of the mouthparts in Palaeodictyoptera: the maxillary stylets were located under the man- dibular stylets (in fig. 7 reversed because of the ventral view of the beak) ; they were distinctly shorter than the mandibles; they were



================================================================================

Figure 8. Blockdiagram representing section of the beak between A and B
(see fig. 71, in ventral view. Left part (Md, indicated by bracket) is probably the natural inner surface of the mandibular stylet where it contacts maxillary stylet. Right part is a composite of several inner sur- faces of the hollow mandible. Oval elevations (E) and crescent-shaped pits (P) are probably opposite ends of short perpendicular pillarsl which crossed the mandibular cavity and supported it from inside. Original. Lower Permian of Czechoslovakia.
divided into more robust galea and thin, protruding Iacina (fig. 7-L) ; the lacinia is located at the inner margin and underneath the galea; the Iacinia extends beyond the beak and has two inward curved apical lobes; the external surface of the maxillae carried fine ridgesO2 Because the beak is split along the median plane, it presents an uniquely favorable occasion to study the inner structure. However, it gives little reliable information about the character of the food canals. The transverse section of the beak (fig. 8) is actually a com- posite of several inner surfaces of the mandibles, all of them carrying alternating grooves and ridges, and is slightly distorted by an oblique pressure. Because of the uneven level of splitting and slight defoma- tion during fossilization, the food canals are not clearly distinguish- able.
The maxillary palpi are robust and overlap with the beak (fig. 6). All segments carry a flat-topped longitudinal ridge, The surface is
covered with a fine rugosity and with occasional irregular grooves. 'The more perfect preservation of the beak in MonsteroPtemm morawi- cum brings an explanation of the ((protruding needle-like tips" in the beak of Mecynostoma dohrni (Palaeodicty~ptera~ Mecynostomatidae, Kukalovi 1969, p. 2101 fig. 28). The protruding structures are undoubtedly also laciniae extending beyond the superimposed pair of mandibles.



================================================================================

Psyche [September
Figure 9. Monsteropferum moravicum n.sp. Enlarged ovipositor with striations, scattered setae and stylus (S), resembling the endophytic ovi- positor of Odonata. Lower Permian of Czechoslovakia. The legs expose their natural ventral surface. The coxae are short and conically truncate (fig. 6-C; fig. IoJC). Proximally are ad- joined two additional circular structures, which perhaps represent the katapleuran ring separated by a sigmoidal paracostal suture from the anapleuran ring (fig. 10-Ky S, A). The trochanter (fig. 6-T; fig. IO- T) is grown together with the femur and is preserved only as a triangular swelling. The tibia is slightly longer than the femur. The tarsus is five-segmented, with the 3rd ,and 4th segments distinctly shorter than the remaining ones. The praetarsus bears a pair of lat- eral claws, The legs are covered by scattered setae and a granular rugosity .
The mesothorax and metathorax are about equal in size) with a broad V-shaped ridge.
The ovipositor has striations (fig. 9) similar to those in some other Palaeodictyoptera (Kukalovi in Carpenter, I 97 I ) p. I 241, fig. 6) and in the related order Diaphanopterodea (Kukdovi 1961, p. 293, fig. 2). The ovipositor carries scattered, proximally oriented, setae. With this specimen styli are recognized for the first time for the order Palaeodictyoptera. Within the extant insects) styli on female genitalia are known in the adult stage only in one order, the Ode nata, in which they arise from the second coxopodite. Also, the gen- eral appearance of the ovipositor is very much like the endophytic ovipositors of some Odonata.
The projections (fig. 6-P) are long and very branched, apparently much more than preserved in the fossil, They have scattered setae and their surface is finely rugose. The sockets of the setae are deeply incised and their density is about equal to that on the legs. The setae increase in number towards the ends of the branches. The projections are stiffly backwardly curved and were undoubtedly sclerotized. Previously, I have been able to study projections in four families of the order Megasecoptera. In all the projections were simple, un-



================================================================================

Kukalovd-Peck - Palaeozoic Insect order^ 265
Figure 10. ,Monsteropterum morawicum n.sp. Enlarged proximal parts of left front leg and left middle leg. A = anapleuran ring; C = coxa; f = front leg; K = katapleuran ring; m = middle leg; S = sigmoidal paracostal suture; T = trochanter. Original, ventral view. Lower Per- mian of Czechoslovakia.
branched outgrowth of the terga. The specimen here described is the first example with projections in the order Palaeodictyoptera and at the same time the first one to show branching. Additional exampk should be expected within Palaeodicty~ptera~ even though their oc- currence probably was not as common as in the Megasecoptera. Dimensions: Beak length 20 mm, width at the middle 2 mm; lacinia length I mm; complete segment of maxillary palpus length 8 rnm; legs I. pair: coxa length 2.8 mm, femur length 10.2 mm, tibia length 12.2 mm, tarsus length 8.2 mm, praetarsus length 2.6 mrn ; 11. pair : coxa length 2.4 mm, femur length 12 mm ; mesothorax length 4.8 mm, width 14.6 mm; metathorax length 4.8 mm, width 14.6 mm ; abdomen total length about 38 mm ; fragment of ovipositor length 9.7 mm; longest fragment of abdominal projection length 16 mm; maximum width 0.8 mm.




================================================================================

266 Psyche
[September
Original: No. 5/1g72 (obverse and reverse of the body and basal parts of wings; separately reverse of the right front leg) ; Paleonto- logical Institute of Charles University, Prague, Czechoslovakia. Col- lected in the Lower Permian deposits near Obora, Moravia. SUMMARY
Megasecoptera and Palaeodictyptera, two of the three extinct paleopterous haustellate insect orders related to extant Ephemeroptera and Odonata, were found to carry fringe-like ~rojections on the thorax and abdomen. Both orders include mostly large to very large insects with good flying ability, which held the wings outstretched when resting. The nymphs were terrestrial, probably arboreal, and had articulated wing pads which were oriented obliquely backwards (Carpenter and Richardson, I 968).
The projections are hollow outgrowth of the terga, forming usually regular rows along the posterior margin or occasionally also on the nota. They are more or less sclerotized, protruding up and back- wards from the body. The surface is rugose and with scattered tac- tile setae. The projections are simple or richly branched and vary from short to very long. They occur in both adults and nymphs. As far is known, their morphology is characteristic for separate fam- ilies.
Now, the projections are known in Mischopteridae, Aspidothora- cidae, Corydaloididae, Brodiidae, Protohymenidae and Bardohy- menidae in the order Megasecoptera and in H~~moiopteridae in the order Palaeodictyoptera. In their location, they are homologous with the dentation on the tr,ansverse abdominal carinae, present in a11 ex- tant Odonata and with paired pits on abdominal terga in nymphs 'of Gomphidae.
Since the projections are obviously a unique and isolated character, their function in two extinct Paleozoic orders is obscure. From the morphology it might be assumed that they were at least to some ex- tent movable and were provided with mechanical sense organs. The following additional characters have been added to the knowl- edge of Palaeodictyoptera :
The beak consists of stylets of mandibles and maxillae, the pair of mandibles being longer and superimposed. Each maxilla is divided into a robust galea and thin lacinia, located at the inner margin and underneath the galea. The lacinia has ,two, inwardly curving apical lobes and extends beyond the beak. Mandibular stylets partly overlap each other along the inner margin.
On contact with the maxillae,




================================================================================

19721 KukaZov&Pec& - PaZaeozoic Insect Order$ 267 they have alternating ridges and grooves, enabling firmer connection. The mandibular stylets are hollow and the cavity is crossed by rows of perpendicularly oriented pillars, providing additional mechanical support.
The legs have short, conically truncate coxae. Proximally, on the ventral side) there are two circular structures, which perhaps rep- resent the katapleuran ring separated by a sigmoidal paracostal su- ture from the anapleuran ring. The trochanter is triangular, grown together with the femur as in Odonata.
The ovipositor is provided by styli as in extant Odonata. LITERATURE CITED
BRONGNIART, C.
1885. Les insectes fossiles des terrains primaires. Bull, SOC. Rouen: - 50-68.
1885. Die fossilen Insekten der primaren Schichten. Jahrb. k.k. Geol. Reichsanstalt) 3 5 (4) : 649-662.
1890. Note sur quelques insectes fossiles du terrain houiller qui pri- sentent au prothorax des appendices aliformes. Bull. SOC. Philo- matique de Paris) 8 ser. 2 (3) : 154.
1893. Recherches pour servir A l'histoire de insectes fossiles des temps primaires. 493 pp. Atlas) 44 pp.
BRAUER) F.
1886. Ansichten uber die palaozoischen Insekten und deren Deutung. Ann. k.k. Nat. Hofmus., Wien) 1: 87-125. CARPENTER, F. M.
1930, The Lower Permian Insects of Kansas. Part 3. The Protohy- menoptera, Psyche) 3 7 (4) : 343-374.
1933. The Lower Permian Insects of Kansas. Part 6. Delopteridae) Protelytroptera) Plectoptera and a New Collection of Proto- donata) Odonata, Megasecoptera) Homoptera and Psocoptera. Proc. Amer. Acad. of Arts & Sci., 68 (11) : 411-503. 1947. Lower Permian Insects from Oklahoma, Part 1. Introduction and the Orders Megasecoptera, Protodonata, and Odonata. Proc. Amer. Acad. of Arts & Sci.) 76(2) : 25-54. 1951. Studies on Carboniferous Insects from Commentry) France. Part 11. The Megasecoptera. Jour. of Paleont.) 25(3) : 336-355. 1962. A Permian Megasecopteron from Texas. Psychej 69 (1) : 37-41. 1971. Adaptation among Paleozoic Insects. Proc. North Amer. Paleont. Convention, Sept. 1969. Part 1: 1236-1251. CARPENTER, F. M. AND E. S. RICHARDSON, JR. 1969. Megasecopterous Nymph in Pennsylvanian Concretions from Illinois. Psyche) 75 (4) : 295-309.
1971. Additional Insects in Pennsylvanian Concretions from Illinois Psyche) 78 (4) : 267-295.
CRAMPTON, G. C,
1927. Eugereon and the ancestry of the Hemiptera, psocids, and Hymenoptera. Bull. Brook. Ent. SOC. 22: 1-17.



================================================================================

268 Psyche [September
HANDLIRSCH, A.
1906. Die fossilen Insekten und die Phylogenie der rezenten Formcn. Leipzig. 430 pp.
KUKALOV.~, J.
1961. Palaozoische Insekten in der Tschechoslowakei. Proc. XI. Int. Entomol. Congress : 290-294.
1969. Revisional Study of the Order Palaeodictyoptera in the Upper Carboniferous Shales of Commentry, France. Part 1. Psyche, 76 (2) : 163-215 ; Part 2. Psyche, 76 (4) : 439-486 ; Part 3. Psyche, 77 (1) : (1970) : 1-44.
LAMEERE, A.
1908. La palkontologie et les mktamorphoses des insectes. Ann. Soc. Ent. Belgique, 52 : 127-149.
1917. Note sur les insectes houillers de Commentry. Bull. Soc. 2001. Fr., 42: 27-37.
1917. Revision sommaire des insectes fossiles du Stephanien de Com- mentry. Bull. Mus. Paris, 23: 141-200.
LAURENTIAUX, D.
1952. Dkcouverte d'un rostre chez Stenodictya lobata Brgt. (Palkodic- tyoptiire stknodictyide) et le probliime des Protohkmiptiires. Bull. SOC. G6ol. Fr., skr. 6, 2: 233-247.
1953. Classes des Insectes. In Traitk de Paleontologic (Piveteau), Paris. 3 : 415.
MARTYNOV, A. V.
1938. Etudes sur l'histoire gkologique et de phylogenie des ordres des insectes (Pterygota). Ie partie. Trav. Inst. Paleont. Acad. Sci. U.R.S.S., 7 (4) : 1-150.
1941. Permian fossil Insects from Tshekarda. Trav. Inst. Paleont. Acad. Sci. U.R.S.S., 11 (1) : 5-62.
NEEDHAM, J. G., M. J. WESTFALL, JR.
1955.
A Manual of the Dragonflies of North America (Anisoptera) : 615 pp.
TILLYARD, R. J.
1924. Kansas Permian Insects. Part 3. The New Order Protohymeno- ptera. Amer. Journ. Sci. (5), 8 (44) : 111-112. WALKER, E. M.
1953. The Odonata of Canada and Alaska. I.: 292 pp. ZALESSKY, G.
1937. Etudes des insectes permiens du bassin de la Sylva et prob- lkmes de I'kvolution dans la classe des insectes. 1. Sur un nouveau representant des Protohymenoptkres et sur les voies du proces d'kvolution dans la morphologie de la nervation des ailes de ce groupe. Problemy paleont., 2-3 : 601-607.



================================================================================