F. M. Carpenter and Eugene S. Richardson, Jr.
Additional Insects in Pennsylvanian Concretions from Illinois.
Psyche 78:267-295, 1971.

Full text (searchable PDF, 9692K)
Durable link: http://psyche.entclub.org/78/78-267.html

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

ADDITIONAL INSECTS IN
PENNSYLVANIAN CONCRETIONS FROM ILLINOIS
The ironstone nodules from the Francis Creek Shale (Mid- dle Pennsylvanian) of Illinois continue to yield many interesting and significant insects. The specimens described in this paper were obtained in former mine pits in Grundy, Will and Kankakee Counties3, and have been made available to us by the following col- lectors, who have been unusually successful in finding insects: Mr. Jerry Herdina, Berwyn, Illinois; Mr. Joseph Makowski Chicago; Helen and Ted Piecko, Chicago; Mr. Paul Tidd, Mendota, II- linois; and Mr. and Mrs. Francis Wolff, Park Forest, Illinois. We are most grateful to them for their cooperation in loaning their specimens to us for study and their patience in waiting for the re- results. Special thanks are extended to Mr. Jerry Herdina and to Helen and Ted Piecko for allowing us to photograph and to make a thorough examination of all the insects in their collections. Sub- sequent papers in this series will deal with additional specimens which they and other local collectors have fo~nd.~ The insects discussed in this paper belong to four orders: Palae- odictyoptera, Megasecoptera, Prodonata and Protorthoptera. All of these specimens are of unusual interest for one reason or another, but two of the specimens in Mr. Herdina's collection are of excep- tional significance; one is the first unquestioned nymph of the order Palaeodictyoptera that has been found and the other is a brachy- pterous adult of a protorthopteron.
ORDER PALAEODICTYOPTERA
Family Lycocercidae Handlirsch
Among the Palaeodictyoptera in the collections at hand there are two species referable to the family Lycocercidae. One of these is in the Herdina collection and the other is in the Field Museum col- lection.
- -
'Harvard University, Cambridge, Mass. 02138 'Field Museum of Natural History, Chicago, Illinois 60605 'Pit Six is about three miles northeast of Coal City, about on the Will- Grundy County line; Pit Eleven is about three miles south of Braidwood, on the Will-Kankakee County line, in northern Illinois. 'Partial financial support of this research is gratefully acknowledged to the National Science Foundation: Grant No. GB27333, F. M. Carpenter, Harvard University, principal investigator; and Grant No. GB8266 R. G. Johnson, University of Chicago, and E. S. Richardson, Jr., Field Museum of Natural History, principal investigators.



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

268 Psyche [December
The Herdina specimen is a nymph with its wing pads held ob- liquely away from the body (figure I). In this respect the wing pads resemble those of the megasecopterous nymph, Mischoptera doug- lassi, described from the same formation a few years ago (Carpenter and Richardson, 1968). However, the venation in the present nymph is sufficiently well indicated in one of the wing pads to show that the insect is a member of the Palaeodictyoptera and that it prob- ably belongs to the family Lycocercidae. This is of unusual signifi- cance, since all of the nymphs which have previously been assigned to the Palaeodictyoptera actually belong elsewhere or have very du- bious palaeodictyopterous affinities. Although until recently the family Lycocercidae has been known only from the Upper Car- boniferous of Europe, it has lately been found in Pennsylvanian de- posits of New Mexico ( Carpenter, 197 I ) . Genus Lycodemas, new genus5
It seems advisable to refer this nymphal form to a separate genus rather than to assign it to one of the three genera of the family al- ready recognized (Kukalovi, 1969) ; the venation of the nymphal wing is not fully developed and does not provide a satisfactory con- cept of the adult venational pattern for comparison with the other known lycocercid species. Since the posterior media (MP) is much less developed in the nymph than it is in the known adults of Lyco- cercus and the related genus Apopappus, we consider that this (i.e., the less developed MP) should be the diagnostic feature of the genus Lycodemas. The more obvious peculiarities of the nymphal wing, such as the narrow proximal region, are aspects of the immature state of its development.
Type species : Lycodms adolescens, 11. sp. Lycodemas adolescens, n.sp.
Figures 1-3
Length of fore wing pad, I I mm.; width, 3 mm. ; length of body from front of mesothorax to end of abdomen (as preserved), 26 mm. ; width of first abdominal segment, 6 mm. The venation, as faintly indicated in a fore wing pad, is represented in figure 2. The wing sheath (w) is conspicuous around the wing except in the ap- ical region, where it is narrow. The subcosta (SC), which extends nearly to the apex of the wing, has a series of blunt projections or tubercles near the middle of the wing; these may possibly have been setal bases in the living insect. The precise origin of Rs from R is 'The generic name is derived from a combination of lykos (wolf) and dmas (body) and is considered neuter.




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

19711 Carpenter ‰â Richardson - Pennsylvantun Insects 369 Figure 1. Lyrodemar adolestem, n.sp. Photograph of hoiotype, No. H413, Herdina collection.




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

Psyche
[December
Figure 2. Lycodemas adolescens, nsp. Drawing of holotype. W, wing sheath; other abbreviations are the conventional venational symbols. not visible in the wing pad but it presumably arises near the basal part of the wing, as indicated in the drawing. Rs has ten primary branches, with one or two secondary branches; the area between MA and the preserved part of MP presumably had some anterior branches from MP but they are not discernible in the specimen; in any event, MP seems to be less developed than in other genera of this family. CUP is extensively branched and conforms to the usual pattern in the Lycocercidae. The narrow form of the wing basally is probably due to the immature nature of the wing pad. Cross veins are discernible in a few small areas; they are apparently numerous and reticulate. The general venational pattern, as far as it can be determined, is shown in figure 2.6
The venation of the hind wing appears to be very similar to that of the fore wing. The wing pads on one side of the body seem some- what broader in the photograph (figure I) than the other pair, but this is deceptive; the matrix was chipped away for some distance from these wing pads, giving the impression that the exposed area was actually part of the wing. The wing pads on both sides are 3 mm. wide.
Holotype:
No. H413a, b; collected in Pit Eleven; in the collec- tion of Mr. Jerry Herdina, Berwyn, Illinois. The most interesting feature of this nymph is the position of the wings with respect to the body.
The arrangement is very similar
indeed to that which we have described in the megasecopterous nymph, Mischo@tera douglassi Carpenter and Richardson. As in the latter, the wing pads of Lycodemas have no contact with the body 'We are indebted to Dr. Jarmila Kukalova-Peck for the preparation of this figure.




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

1971 Carpenter ‰â Richardson - Pennsylvanian Insects 27 I Figure 3. Lycodemas cf. adolescens. Drawing of specimen No. H110, Herdina collection.
except in the articular area of the wings themselves, in contrast to the structure of the wing pads in the nymphs of Recent insects, in- cluding those of the Ephemeroptera. A discussion of the evolution- ary significance of this position of the wing pads is contained in our previous paper cited above and in Dr. Kukalovi's account of the Permian may-fly nymphs ( I 968).
The type of L. adolescens is the first nymph known that can with- out question be assigned to the Palaeodictyoptera; the nature of the venation in the wing pad seems to provide conclusive evidence of the palaeodictyopterous nature of the insect, especially in view of the homonomous condition of the wings. It is regrettable, of course, that more of the body structure is not preserved. What little is visible gives no indication that the nymph was modified for an aquatic existence; this is consistent with the more extensive evidence pro- vided by the nymphs of Mischoptera for the Megasecoptera. In the Herdina collection there is also a single wing (No. HI 10, Pit Eleven) ; it is obviously a nymphal wing, since it is included in a sheath (figure 3). It has a length of 27 mm. and a maximum width of 7.5 mm. and is therefore three times the size of adolescens. In all probability, although no proof can be given, this fossil is an older nymph of adolescens or of another species of the genus. The wing has the slightly falcate shape that is present in the mature wings of the Lycocercidae (see Kukalovi, 1968, figs 33 and 34). The specimen in the Field Museum collection is being assigned tentatively to the Lycocercidae. Assignment to this family i,s based on what little is preserved of the venation of one wing, probably about half the entire wing; MP is extensively branched, much more so than in the Dictyoneuridae; CuA is unbranched but CUP is well developed; the anal veins are apparently numerous. On the basis of the wings alone, especially in view of their fragmentary na- ture, this insect would hardly warrant formal description and nam-



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

Psyche
ant /
[December
Figure 4. Notorachis wolfforum, n.sp. Drawing of holotype, No. PE 21699, Field Museum; ant, antenna; hind wing represented by dotted lines. Black circles on abdomen mark the locations of spines on the posterior edges of the tergites; the probable lateral margins of the abdomen are indicated by broken lines.
ing.
However, the prothoracic lobes of this species are extraos-di- narily modified, nothing comparable to them having previously been observed in the Palaeodictyoptera. Since this insect is of unusual interest, generic and specific names are being assigned. Genus Notorachis, new genus"
This genus is apparently related to Lycocercus. The outer mar- gins of the pronotal lobes are heavily sclerotized and form a series of long spines; the more basal portion of each lobe is less sclerotized but there is a double row of blunt setae or tubercles extending trans- versely across each lobe (preserved as pits in the obverse). In Lyoo- 'The generic name is derived from a combination of noton (back) and rachis (spine) and is considered feminine.



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

19711 Carpenter ^5 Richardson - Pennsylvanian Insects 273 cercus, although the lobes are prominent, they are membranous or nearly so with a radiating series of vein-like structures, as in most other Palaeodictyoptera; paranotal spines are unknown in Lycocercus. Type species: Notorachis woZff'oru/n, nsp. Notorachis wolff orurn, n.sp.
Figures 4-6
Length of wings, as preserved, 22 mm.; maximuum width of fore wing, 10 mm.; width across pronotal lobes, including spines, 20 mm.
The
fossil consists of the dorsal aspect of the head, pronoturn, and meso- and metathorax; a few segments of the abdomen are in- dicated and the wings are outstretched in the palaeopterous ~osition. The fore and hind wings on each side overlap but the basal part of the venation of one wing is discernible for the most part. The costal space seems to be of uniform width; the precise origin of the radial sector is not preserved; MA obviously arises before the ori- gin of the radial sector and is roba ably unbranched, although its distal part is not preserved; MP has at least five terminal branches and Cup is even more extensively branched. The hind wing is almost completely covered by the fore wing and only a few veins can be distinguished.
The most conspicuous of the body structures are the pronotal lobes (figures 5 and 6), which are not sharply set off from the rest of the prothorax. The margins of the lobes are heavily sclerotized and bear seven prominent spines on each side, the first and last of these being somewhat shorter than the others. The two posterior spines of each side are directed somewhat do'rsally as well as laterally. Extending along the median axis of each lobe is a double row of setae (preserved in the reverse of the fossil as fine pits). The sur- face of the slightly elevated margin of the pronotal lobe is smooth along the posterior edge of the lobe; on the anterior edge it is occu- pied by scattered setae or fine tubercles. Similar setae are present on the proximal third of all the spines. Somewhat before mid-length of each spine the setae are smaller and directed posteriorly; in the distal potion of each spine, the setae are succeeded by strong, longitudinal ridges.
The head is visible just anterior to the lobes but there is no sharp separation in the fossil between the head and thorax; two relatively large, circular structures are visible on the sides of the head, pre- sumably the compound eyes. A four-millimeter portion of one an- tenna is preserved; it appears to arise from above the center of the



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

Figure 5. Notorachif wal^orum, n.sp. Photograph of holotype (reverse), showing overlapping of wings and die general form of the body. eye and curves laterally. Its proximal segments are less than half as long as wide; the distal segments are relatively longer and are clad with fine prostrate setae. The preserved part of the antenna is surprisingly thin and delicate.
The meso- and metachorax seem to form a compact unit, each of the segments being relatively short. Portions of two segments, prob- ably tibia and tarsus, of a weakly sclerotized fore leg, are preserved anterior to one of the pronotal lobes. The portion of the tibia is about three rniiiirneters long and the tarsus nearly two millimeters, The tibia is armed with short spines directed distally, and the tarsus with setae. There is no reason to suppose that this leg was adapted for anything but slow walking.
P .
1 he abdomen is indicated by a series of faint, transverse depres- sions that mark the segmentation. It is preserved for a length of



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

19711 Carpenter &f Richardson - Pennsylvanian Insects 275 about 23 mm., to the edge of the nodule, and it undoubtedly orig- inally extended beyond that point. Pairs of strong, flat, setiferous spines project dorsally and somewhat posteriorly from the abdomen, arising directly on the posterior margins of the tergites. The first spine is short; the second, third and fourth are nearly as long as the width between them; the fifth is short; a very slight projection marks the position of the sixth and there are none in the more posterior positions. These spines do not appear to mark the lateral edges of the tergites, which do not have their full widths preserved. The spines that are preserved in the fossil are about 3.5 mm. apart laterally and were almost certainly a little to each side of the median longitudinal axis of the abdomen. Other spines along the posterior tergal margins were probably present near the sides but can only be assumed since the lateral portions of the segments are broken away. In all probability the tergal spines in Notorachis were similar to those already described in the megasecopterous Mischoptera douglassi, though the strong, dorsally projecting spines just described are flat- tened parallel to the body axis rather than to the surface of the tergites. (See Carpenter and Richardson, 1968, p. 306). It is pertinent to note that prothoracic spines were also present in Mis- choptera.
Holotype : No. PE2 I 699, Field Museum of Natural History; from the collection of Mr. and Mrs. Francis A. Wolff, Park Forest, Illinois, who collected the unique specimen in Pit Eleven. The spe- cies is named for both Mr. and Mrs. Wolff. This insect is unusual, among the Palaeodictyoptera so far known, in having elaborately modified pronotal lobes; the only other species that may approach Notorachis wolfforum in this respect is Stiibocro- cis heeri (Goldenberg) (Upper Carboniferous of Germany), which is, however, a member of the Dictyoneuridae. In Stilbocrocis the lobes are smaller than in Notorachis and possess fewer spine^.^ The most puzzling aspect of this fossil is the shortness of the pterothorax and the consequent overlapping of the fore and hind wings. Since this overlapping is symmetrical for both pairs of wings, it is almost certainly not due to distortion during preservation. Fur- thermore, the thorax itself is so short that the overlapping would 8~tilbocrocis has consistently been represented in the literature as having large pronotal lobes, bearing a series of radiating veins within the lobes themselves. From a recent examination of the type specimen (on deposit in the Natural History Museum at Bonn), I am convinced that the radiat- ing ridges are actually spines which project beyond the edges of the lobes. A detailed account of this fossil will be published elsewhere. (FMC).



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

276 Psyche [December




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

19711 Carpenter &f Richardson - Pennsylvanian Insects 277 Figure 7. Homalonevra dtibasindasi Carpenter. Photograph of speci- men in Tidd collection. The arrow points to the cross-section of the beak. seem to have been unavoidable in the living insect. On the other hand, such an arrangement of the wings would seem to be very in- efficient mechanically. It is in this connection interesting to note that the only other palaeodictyopteron known that shows a similar overlapping of the fore and hind wings and a comparable, short thorax, is Stilbocrocis heeri, already mentioned. This condition of the wings is readily seen in the photograph of the specimen given by Guthorl (1934), as well as in the figures by Goldenberg (1854), Schlechtendal ( 191 2) and Guthorl ( 1934) ; it is not shown in Handlirsch's highly imaginative reconstruction of the insect ( 1920) in which the wings are represented in normal position. As noted above, fiilbocrocis has the wing venation characteristic of the Dic- tyoneuridae, whereas Notorachis has a very different venational pat- tern, allying it to the Lycocercidae.




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

Psyche
[December
Figure 8. Homaloneura dabasinskasi Carpenter. Drawing of specimen in Tidd collection.
Family Spilapteridae Brongniart
Homaloneura dabasinskasi Carpenter
Figures 7-9
Ho~naloneura dabasinskasi Carpenter, I 964 : I I 7 A second specimen of this species, also in an ironstone concretion from the Francis 'Creek Shale, was collected in 1971, by Mr. Paul Tidd of Mendota, Illinois, in Pit Eleven and is now contained in Mr. Tidd's collection (figure 7) It consists of the basal part of a fore wing, portions of the head and thorax and traces of a hind wing. Determination of the species is based on the fore wing; except for minor points, such as a deeper fork on CUP, the venation is like that of the holotype. The wing markings, characteristic of the spe- cies, are faintly preserved. Some of the thoracic structures are of interest, since they are known in only a few genera of the order. The thoracic segments are unequal (figure 8) the prothorax being slightly the smallest of them. There are no indications of the prothoracic lobes, which have previously been described in Homa- loneura (Kukalovi, 1969). The fore legs are preserved in the ob- verse half of the specimen; they are robust and possess numerous



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

19711 Carpenter & RichardsonÌÔPennsylvania Insects 279 Figure 9. Homdonfvra dahstliskasi Ciirpmter. Photograph of cross- section of beak; specimen in Tidd collection. The arrow points to the median stylet.
setae, as in other Falaeodlctyoptera. The coxae are prominent and in the obverse half of the fossil extend anteriorly from the pro- thorax. The tarsal segmentation is not discernible, except for the first tarsomere.
The head is only vaguely indicated in the fossil (reverse half) but just anterior to the head itself there is a transverse section of the sucking beak, this being visible at the point- where the two halves of the concretion separated. In order to appreciate the sig- nificance of this section, it must be borne in mind that conclusive evidence has now been provided, in the publications of Crampton (1927), Lameere (1933), Laurentiaux ( 1952 and 1953) and Ku- kalovi ( 1969 and 19701, showing that all of the Palaeodictyop- tera possessed a haustellate beak. According to observations pre- viously made (and summarized by Kukalovi, igp), the beak consisted of four slender stylets, apparently supported ventrally by a long and somewhat broader labium, usually transversely ridged.



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

280 Psyche [December
The beak was not directed ventrally from the head but anteriorly and ventrally.
The section of the beak in the new specimen of dabasinskasi is about 3 mm. in front of the anterior margin of the head (figures 7 and 8) ; it presumably shows the structure of the beak close to the head, not at a point further along the beak. As shown in the photo- graph (figure g), there is a vague circular area of discoloration which appears to mark off the area of the beak. Within this can be seen sections of two pairs of stylets; these are very dark and distinct. The anterior pair, which are somewhat the larger, are presumably the mandibles, and the other pair, the maxillae. These four stylets are symmetrically arranged within the beak and with respect to each other. However, in addition to these there is a fifth stylet, midway between the maxillae and slightly more posterior. The sym-
metry of the pattern formed by these five structures, as seen in sec- tion, is very striking. It is virtually certain that the fifth stylet is one of the mouthparts, presumably derived from the hypopharynx; the latter, of course, is a median structure and it does function as one of the stylets of the haustellate mouth-parts of some Diptera and Hemiptera. The labium, which as noted above is broader and thicker than the stylets in the Palae~dict~optera, is missing from the section, as are the large maxillary palpi. In any event, it now seems clear that in the spilapterid Palaeodictyoptera, at least, there were five stylets in the beaks, with an arrangement which closely parallels that in some Recent insects.
Genus Spilaptera Brongniart
In the Herdina collection there are two spilapterids that appear to belong to the same species.
Only two representatives of this
family have been previously reported from the Francis Creek Shale, or, in fact, from all Pennsylvanian deposits in North America: Homaloneura dabasinskasi Carpenter and Mcluckiepteron luciae Richardson. The new specimens belong to an undescribed species, which we are assigning to Spilaptera, otherwise known only from the Con~mentry shales of France.
Spilaptera differs from Homaloneura in having several long, sig- moidal and oblique cross veins between RI and Rs, beyond mid- wing. In Homaloneura these cross veins are straight or nearly so and they are transverse, or only slightly oblique. CUP is usually unbranched in Hoinalioneura, although it may be forked distally, very near its termination. In Spilaptera CUP ranges from unbranched to deeply forked. The new species described below has the sigmoidal



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

19711 Carpenter 8 Richardson - Pennsylvania?! Insects 281 Figure 10.
463, Herdina
Spilattera americana, n.sp. Drawing of holotype, No. H collection. Length of wing as preserved, 38 mm. a,nd oblique cross veins present and it is accordingly placed in the genus Spil~ptera. The several other known genera of Pennsylvanian Spilapteridae, including Mcluckiepteron, have very distinctive fea- tures, such as numerous cross veins, more extensive branching of veins, pronounced tapering of wing, etc. Spilaptera americana, n.sp.
Figure 10
Length of wing (as preserved) 38 mm.; maximum width, 15 mm. Costal margin distinctly concave, slightly more so than in the type species of the genus, S. packardi; Sc extending to wing apex; Rs with 6 terminal branches, none forked in types; MA with 4 branches, CuA with 5; 'CUP deeply forked, almost to its origin. Cross veins not numerous, about as many as in S. packardi, forming poorly de- fined rows. The sigmoidal cross veins between RI and Rs are long and more oblique than in 8. packardi. The wing markings consist of