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PENNSYLVANIAN INVERTEBRATES
OF THE Mazon Creek Area, Illinois

INSECTS

EUGENE S. RICHARDSON, JR.
Curator of Fossil Invertebrates

FIELDIANA: GEOLOGY

VOLUME 12, NUMBER 2

Published by

CHICAGO NATURAL HISTORY MUSEUM

JANUARY 25, 1956

PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS

Insects

INTRODUCTION

Pennsylvanian insects have been found in only three locaHties
in IlHnois. The most important of these is the area in Will and
Grundy counties that comprises the natural outcrop of the Francis
Creek shale member of the Carbondale Formation on Mazon Creek,
and also the artificial exposure nearby, created by strip mining for
coal (see no. 1, this volume). One Pennsylvanian insect is known
from Colchester, McDonough County, and five from Vermilion
County, south of Danville; in contrast, 130 species have been
described from Mazon Creek and the related strip-mine exposures,
and seven more are included in this paper.

Descriptions of the first two species of Pennsylvanian insects to
be described from the Mazon Creek sites were published in 1864 by
J. D. Dana. S. H. Scudder described 36 more, including the species
from Colchester and one from the Danville area. Dana and Scudder
represent what has been termed the "Primitive Period" of fossil
insect work in Illinois (Richardson, 1953), a period that closed with
a paper by A. L. Melander in 1903. The extinct orders were not
then understood; fossils were assigned to broadly conceived modern
orders and were compared with modern genera or were placed in
artificial groups in the belief that insect orders had not become
differentiated by Pennsylvanian time.

The "Middle Period" of Illinois paleoentomology, to pursue the
figure, is dominated by Anton Handlirsch, of Vienna. From 1906
to 1920, Handlirsch described 77 Illinois species, almost all from
Mazon Creek. Having a great many more species for comparison,
Handlirsch had a better insight than had Scudder or Dana into the
problem of a natural classification. He named four new extinct
orders for the Pennsylvanian insects and redefined others, giving
the classification of Paleozoic insects a validity potentially equal to
that of modern insects. His observations and his taxonomy have
not withstood all tests of later work, but his papers are still a neces-
sary starting point for any one working with Mazon Creek insects.

15

16 FIELDIANA: GEOLOGY, VOLUME 12

Balancing the Primitive and Middle periods there is inevitably
a Modern Period, which is based on the work done by R. J. Tillyard,
F. M. Carpenter, and A. V. Martynov on the Permian insects of
Australia, Kansas, and Russia. Attention was for a time turned
away from Illinois fossil insects, largely because few new specimens
were being found; but later, F. M. Carpenter (1938, 1943b) described
six species, one from Mazon Creek and the rest from the strip mines.

Since insects are very rare in the Francis Creek concretions, any
significant rate of discovery of new specimens depends on a high
rate of collecting. In the years 1937-46, Mr. George Langford, then
engaged in business in Joliet, Illinois, spent the astounding total of
1,100 days in the field. His collections of that period are now
deposited in the Illinois State Museum, Springfield; the Peabody
Museum, Yale University; and the St. Paul Academy of Science,
St. Paul, Minnesota. His insects are all in the collection at Spring-
field, and the finest ones have been described (Carpenter, 1943b).
Since he joined the staff of Chicago Natural History Museum in
1946, Mr. Langford has been collecting for this institution, and to
date he has found a dozen specimens of insects, the best of which are
described below.

Mr. Langford's collection has been rivaled by the numerous
enthusiastic amateur collectors who visit the spoil heaps of the strip
mines; some insect specimens are known to be in their collections.
One of these, included below, is from the collection of Mr. and Mrs.
John M. McLuckie, of Coal City, Illinois. Other specimens, col-
lected by Dr. and Mrs. Robert H. Whitfield, Mr. Jon S. Whitfield,
Mrs. William D. Turnbull, and the writer, are in the Chicago
Natural History Museum collection. The Walker Museum (Uni-
versity of Chicago) and the Chicago Academy of Sciences also have
collections that include insects from Mazon Creek and the strip
mines.

Including the new species described below, the Mazon Creek
insect fauna as now known comprises 137 species. The composition
of the fauna by orders is indicated in the Table, with the faunas of
Great Britain, the Saar basin, and the northern Appalachians
included for comparison. Unfortunately, the Commentry insects
can not be included, as Professor Carpenter's careful and necessary
revision has not yet been completely published. Probably none of
the four faunas shown in the table is sufficiently well known to form
a trustworthy basis for speculation; yet the differences in rank
of the first three orders listed are so pronounced that they should

RICHARDSON: INSECTS 17

prove to be representative of the differences in the complete faunas,
and certain tentative conclusions may be drawn on this assumption.
The Francis Creek fauna lived in a moderately open wood-
land lying little above sea level, near a shore, and being slowly
aggraded by mud and silt deposits— a plant-covered delta plain.
The composition of the insect fauna seems in accordance with this
view. In contrast with the insect faunas of the other areas listed,
the Mazon Creek fauna includes relatively few roaches. If roaches
preferred then, as they do now, a dense and moist forest habitat,
their scarcity suggests a drier, more open forest.

PENNSYLVANIAN INSECT FAUNAS

Ordinal composition of the Mazon Creek insect fauna compared with those
of the British Coal Measures (from Bolton, 1921-22, and Handlirsch, 1922),
the Saar-Lorraine basin (from Waterlot, 1934), and Pennsylvania and West
Virginia (from Handlirsch, 1922, and Carpenter, 1934).

Order Northern Great Saar- West Virginia and

Illinois Britain Lorraine Pennsylvania

% % % %

PalaeodjctypEtera 18 33 55 9

Protorthoptera 57 , v' 9 4 4

Blattaria 21 Z"^'^^ 47 32 84

Protodonata 1 7 — 1

Megasecoptera '...2 1 — —

Caloneurodea 1 — — 1

Minor orders and indet. ... 2 1 9 1

Total species 137 46 56 89

Waterlot (1934, p. 297), commenting on the abundance of
Palaeodictyoptera in the Saar basin deposits, pictured them as
skimming the surface of ponds much as modern dragonflies do.
Thus the Protodonata and the Palaeodictyoptera may have com-
peted for the same ecological niches. The great relative abundance
of Palaeodictyoptera in the Saar basin may have been related to the
absence or scarcity of fish at that place, as Waterlot (1934, p. 298)
has suggested.

On the basis of their having strong walking legs, the Protor-
thoptera have been pictured (Bolton, 1921, p. 7) as spending much
of their life on the ground, possibly along the margins of swamps.
Though their large wings were well adapted for flight, a preference
for ground feeding, if the interpretation is correct, would be in
accordance with the suggestion that the vegetation of the Mazon
Creek delta was more sparse than that of a typical coal swamp.

18 FIELDIANA: GEOLOGY, VOLUME 12

The apparent dearth of Caloneurodea in Europe (excepting at
Commentry) and of Megasecoptera in the northern Appalachians
is probably caused by the slow rate of discovery of fossil insects
rather than by their absence from those faunas. Insects of those
orders and of the Protodonata were undoubtedly less abundant than
were those in the three major orders.

DESCRIPTION OF SPECIES

Order PALAEODICTYOPTERA

The order Palaeodictyoptera has never been adequately defined.
Devised in 1867 by Dohrn (under the preoccupied name of Dicty-
optera) for the remarkable Permian genus Eugereon, the order grew
by accretion until in Scudder's time (Scudder, 1885) it embraced
all the Paleozoic hexapods then known. Subsequent restriction of
the order leaves it now including 36 families with somewhat varied
attributes, but no longer including Eugereon, now placed in the
order Protohemiptera of Handlirsch.

The principle of priority would no doubt require that the ordinal
name Palaeodictyoptera be substituted for Protohemiptera and
another name (or several names) be proposed for the array now
called Palaeodictyoptera. Fortunately, we are not compelled to do
this, though others, notably Cramp ton (1930), have suggested new
ordinal names for portions of the group. By common usage, the
family Stenodictyopteridae (Dictyoneuridae of authors) and its
relatives are considered to constitute the order in question. But
what are the characters of this order? A few that may be listed
are taken from published partial definitions or implied in published
discussions. The following list is concerned only with wing char-
acters; features of the head, thorax, and legs are equally significant,
but manifestly cannot be applied to specimens consisting of wings
alone, as most specimens do.

(1
(2
(3
(4
(5

(6
(7
(8
(9
(10
(11

Longitudinal veins much branched.

MA and MP well developed.

At least 3 anal veins.

Mesh-like archedictyon.

Veins arising independently at base of wing; but anterior and posterior

limbs of dichotomous veins (M and Cu) may arise from a common stem.

Veins do not cross each other or fuse together.

Rs arises in proximal half of wing or about midwing.

Veins curve toward posterior margin.

Anal veins curve toward base of wing.

CuA present, convex.

CuP branched.

RICHARDSON: INSECTS 19

None of these criteria may be accepted without reserve. Tabu-
lation from published figures and descriptions of Palaeodictyoptera
indicates that in some families all reported specimens are so incom-
plete as to preclude use of several of these characters; other specimens
that are complete fail to fulfil one or more conditions.

Just as members of a plexus of orders have an orthopteroid
facies and are commonly placed together in the inclusive order
Protorthoptera, so there is a palaeodictyopteran facies uniting
various species into this primitive order.

In the homologizing of wing veins that is an integral part of the
description of species, the concept of the order Palaeodictyoptera
is in effect a "morphotype" (Zangerl, 1948, p. 355) of the primitive
insect wing. The morphotype of the palaeodictyopteran wing
includes the veins C, Sc, Rl, Rs, MA, MP, CuA, CuP, and wA.
Since within the order the number of branches of these veins and
the number of anal veins are variable, it is fruitless to adopt a numer-
ation for the branches; as Zangerl has shown (op. cit., p. 369),
in cases where undifferentiated structures occur in different quantity
among compared forms in whose mutual morphotype the quantity
is unstable, each of those structures in one form is the homologue
of every similar structure in the compared form. Within a smaller
group, in which, for example, Rs constantly has four branches, the
morphotype of that group will have that specified number of
branches, and corresponding branches in compared forms may be
properly considered homologous. The naming of veins in the
descriptions that follow is limited to the veins that are significant
for that group; where a more detailed nomenclature is employed,
it is for convenience in description and is not intended to imply
a detailed homology.

The Palaeodictyoptera described below pertain to two known
families, one undescribed family, and two undetermined families.

Family Spilapteridae

It is interesting to note the occurrence of a spilapterid in this
fauna. Other species of the family have been reported in Penn-
sylvanian rocks only from France and England; a single Permian
species extends the geographic range to Kansas. Handlirsch (1925,
p. 136) defines the family: "Rs more or less richly branched. Front
branch of M always many-branched, as well as the hind branch,
Cu often with branched anterior member. Anal veins with numer-
ous branches. Straight cross veins more or less widely spaced.

20 FIELDIANA: GEOLOGY, VOLUME 12

Wings often decorated with stripes or spots." Further remarks
on the family are presented below.

Mcluckiepteron, new genus

Description. — Hind wing: Subtriangular in shape, with concave
anterior margin and recurved tip; greatest width near base. Narrow
trichiated fringe of membrane anterior to C. C slender, accom-
panied in proximal half of wing by a broad, thickened, convex ridge
between C and Sc. Sc and Rl very close to C. Sc extending to or
nearly to apex. Rs arising very near base, running parallel to Rl,
converging toward it near apex, sending off a pectinate series of
simple or forked branches in distal half of wing. MA arising near
base from stem of MP, forked at midwing. MP putting forth
a forwardly directed pectinate series of simple branches. CuA
divided near base into two divided branches, their offshoots occu-
pying the triangular space between the main branches. CuP
simple, arising independently. lA branched. Cross veins relatively
few, irregularly spaced, straight, some meeting end to end between
successive veins.

Genotype. — Mcluckiepteron luciae, new species.

Diagnosis. — Mcluckiepteron is allied to Homaloneura and
Homaloneurina of the Commentry fauna and to Dunbaria of the
Kansas Permian. These and perhaps Homaloneurites seem to con-
stitute a compact group within the family Spilapteridae, charac-
terized by the concave anterior margin and long Sc. The new genus
differs from the others in the great development of branches in
CuA, in the pectinate branching of MP, and in the possession of
a precostal fringe.

Remarks. — It is appropriate to name this striking genus in honor
of Mr. and Mrs. John M. McLuckie, of Coal City, Illinois, whose
collection of plant and animal fossils from the strip mines of Will
and Grundy counties contains many outstanding specimens, the
cream of a quarter century's collecting.

Mcluckiepteron luciae, new species
Figures 4, 5

Description. — Hindwing: Length about 55 mm.; greatest width,
at position of termination of lA, 33 mm. Precostal fringe bearing
about three rows of irregularly arranged macro trichia. Auxiliary

RICHARDSON: INSECTS 21

convex coriaceous ridge between C and Sc very broad basally,
narrowing evenly and thinning to nothing at mid wing, where C,
Sc, Rl, and Rs are all bent forward, following the wing margin.
Sc arising in a broadened basal concave sclerotized area about a
third as wide as the convex ridge anterior to it. Rl strong, standing
much higher than C, simple. Rf near base, Rs not branching again
until slightly beyond mid wing; Rs with five or six pectinately
arranged branches, of which the first sends forth two anteriorly
directed twigs. Mf slightly distad of Rf, MA arising as a concave
branch from the concave stem M, but almost immediately becoming
convex and as high as Rl; MA forked slightly distad of the fork of
Rs, the anterior branch continuing the marked elevation of the
stem, but the posterior branch, less elevated, continuing its gentle
curve to the margin. MP with four forwardly directed branches,
the first at midwing, slightly proximad of the fork of Rs. CuAl'
twice forked; CuA2' with three branches, of which the first is forked,
each twig again forking, the second once forked, and the third
simple. CuP distant from CuA and lA, simple. lA at least twice
forked. Cross veins perpendicular or nearly so to longitudinal veins,
tending to slant between veins of different series; sparse except
between Rl and Rs. No trace of mesh-like archedictyon.

Sockets of macrotrichia are to be seen on certain veins of M.
luciae, as follows: on the bottom surface of the main stem of Rs,
but not on its branches; on the top surface of Rl, though possibly
lacking proximad of Rf; probably on the top surface of the basal
portion of CuA. They are plentiful on the thin membranous pre-
costal margin, on both upper and lower surfaces. Wing spots,
appearing as low domes, occupy the distal radial cells.

The broadened convex base of R is split by a corrugation into
two stems that come together just before Rf, the posterior member
of this pair lying tangent to the heavy stem of CuA. The anal area,
as in Permian ephemerids (Tillyard, 1932, p. 107), is incompletely
preserved, probably being formed of a more tenuous membrane
with weaker struts than the rest of the wing. lA, with two forks,
may be seen in that area, as well as part of another anal vein.

Remarks. — Tillyard (1937a, p. 85) described a megasecopteran-
like genus, Kansasia, from the Permian of Kansas, as a spilapterid;
Rohdendorf (1940, p. 108) included it in the order Archodonata.
Handlirsch (1937, p. 116) posthumously proposed the family Dun-

' Designation adopted for convenience of reference, not to indicate a supposed
homology with a similarly named vein in any other wing (see p. 19).

22

FIELDIANA: GEOLOGY, VOLUME 12

bariidae, which he neglected to characterize, for the single genus
Dunbaria; his reasons for separating it from the Spilapteridae, to
which Tillyard (1924, p. 203) had assigned it, are not evident. The
Pennsylvanian Mcluckiepteron and the Permian Dunbaria will be
treated tentatively as members of the same family, though occupy-
ing different branches; they are thus the only spilapterids known

""-^'CuP'.

Fig. 4. Mcluckiepteron luciae, new species. lA, first anal; C, costa; CuAl,
CuA2, first and second forks of anterior cubitus; CuP, posterior cubitus; MA,
anterior media; Mf, fork of media; MP, posterior media; R, radius; Rf, fork of
radius; Rs, radial sector; Sc, subcosta.

from North America, and are, oddly enough, the oldest and the
youngest representatives of an important European family. They
share the character of a thickened basal part of the costal region, and
both have trichia on some of the veins. Whether the other spilap-
terids have these characters cannot be ascertained from Handlirsch's
figures and descriptions. The older genus, Mcluckiepteron, is more
generalized than the younger, in its free CuP, its branched lA, and
its more branched MP, but it is the more specialized in several ways:
(1) it lacks the mesh-like archedictyon ; (2) it has developed the
very unusual precostal membranous margin; (3) Rs is slightly less
branched and includes a forked branch; (4) the auxiliary veins in
CuA are unusually numerous; (5) Mf is situated farther toward the
base; (6) it includes raised wing spots in the radial cells.

The trivial name is given in honor of Mrs. McLuckie, who dis-
covered the holotype and recognized it as a new species of insect
wing. I am grateful to Mr. and Mrs. McLuckie for the loan of the
specimen from their collection.

Fig. 5. Mcluckiepteron luciae, new species, holotype; X 1.7. McLuckie
Collection.

23

24 FIELDIANA: GEOLOGY, VOLUME 12

Holotype. — Two halves of a concretion containing impressions of
a hind wing; in the collection of Mr. and Mrs. John M. McLuckie.
Plaster casts are preserved in the collection of Chicago Natural
History Museum and in the Department of Geology, Princeton
University.

Family Syntonopteridae Handlirsch 1911, emend. Carpenter 1938
Genus Lithoneura Carpenter 1938

Lithoneura Carpenter, 1938, Amer. Jour. Sci., (5), 36, p. 446; 1943, Illinois
State Mus., Sci. Pap., 3, pt. 1, p. 13.

Two species of Lithoneura have been described, L. lameerei
Carpenter 1938 (the genotype) and L. mirifica Carpenter 1943, both
from the strip mines north of Braidwood. The first is known from
both forewings and hindwings, and the second from only a hindwing.
The present species, also from the strip mines near Braidwood, is
known from only a forewing.

Although this species lacks full development of the two charac-
ters prescribed for the forewing in Carpenter's diagnosis of Litho-
neura (Carpenter, 1938, p. 446), the aspect of the wing is certainly
such as to admit it to the genus. Therefore, the generic diagnosis
should be modified to the extent needed to include this species.
True cross veins are only slightly developed in the costal area, being
present in L. carpenteri only in the proximal quarter. The fusion
of Rs and MA is incomplete, these veins being tangent rather than
fused. Cross veins are as well developed in the remainder of the
wing as they are in L. mirifica. The separation of veins normally
fused is not unusual within a genus; in the family Lemmatophoridae,
this character varies within species and even between right and left
wings of an individual (Tillyard, 1928, p. 313; Carpenter, 1935,
p. 107).

Lithoneura carpenteri, new species

Figures 6, 7

Description. — The venation of the forewing is shown in figure 6.
The costa lies slightly inside the margin of the wing basally and
gradually approaches it, so that it is truly marginal only in the
distal quarter of the wing. The narrow precostal area slopes sharply
down from C, forming a flange on the anterior edge of the wing.
Sc extends nearly to the apex, probably terminating on C. Rf is far

— c +

IMPI

Fig. 6. Lithoneura carpenteri, new species; X 2.2. A, anal; C, costa; CuA,
anterior cubitus; CuP, posterior cubitus; I prefixed to vein symbol, interpolated
vein; MA, anterior media; MP, posterior media; MPl, first branch of posterior
media; Rl-3, branches of radius; Rs, radial sector; Sc, subcosta; +, elevated
vein; — , depressed vein.

Fig. 7. Lithoneura carpenteri, new species; X 2.35. Walker Museum no. 45248.

26

26 FIELDIANA: GEOLOGY, VOLUME 12

toward the base; R2+3 separates from R4+5 at about mid wing;
R2 separates from R3 near the apex; R4 + 5 apparently remains
undivided. Intercalary vein IR3 is present. Mf is far toward the
base, but markedly apicad of Rf; MA arises in a strong arch as
a convex vein from the concave stem M and touches Rs but does
not fuse with it; MA bifurcates at Mf, at about mid wing, slightly
apicad of Rf; MP bifurcates about halfway between Mf and Mf,
with IMPl originating at about midwing. CuA arises close to or
from the same stem as M, and bifurcates slightly apicad of Mpf.
CuP, concave, arises independently of CuA, and remains simple,
as in the hind wing of lameerei. Anal area moderately long; anal
veins simple.

In size this species is intermediate between the two previously
known species, and closer to the larger, L. mirifica. The number of
cross veins is likewise intermediate between the two other species.

Remarks. — Since this species is known from only a forewing, and
L. mirifica from only a hindwing, the present specimen might be
thought to represent the unknown forewing of the latter species.
Fortunately, both wings of the genotype are known, so that a test
may be made. A rectangular grid superposed on a drawing of the
hindwing of L. lameerei undergoes a certain distortion in being
transferred to a drawing of the forewing in such a way that inter-
sections of the grid fall on homologous points in the two wings.
If a similar grid transfer is attempted from the hindwing of L.
mirifica to the forewing of L. carpenteri, the distortion is of a dif-
ferent pattern. While this test may not indicate that the wings are
from different species, this is at least implied, and accordingly a new
name is proposed for the species here described.

A curious feature of the wing of L. carpenteri is the presence of
accessory ridges accompanying certain veins, and accessory furrows
accompanying one of the concave veins. Near the base, two slender
ridges appear between R and Rs; both continue parallel and close
to Rl nearly to the apex, where the posterior one ends and the other
blends with Rl. Some, but not all, of the cross veins between Rl
and Rs cross the posterior ridge. Short, shallow furrows are devel-
oped as a bracket around Rf , and a single ridge parallels a portion
of MA before its fork. The accessory ridges resemble those border-
ing the distal portions of the longitudinal veins in Camptoneurites and
Protemhia (Carpenter, 1950, pp. 210-217), Permian protorthop-
terans. Their significance is obscure, but they are probably of no
more than specific or possibly individual importance.

RICHARDSON: INSECTS 27

Intercalary veins, characteristic of syntonopterids, are present
in L. carpenteri in Rs and MP. The convex vein IRS is joined to the
concave R2+3 by a short, weak, concave cross vein. IMPl appears
to originate without connection with MPl.

Holotype. — Two halves of a concretion containing impressions
of the dorsal and ventral surfaces of the wing. Collected in 1938
by Dr. Heinz A. Lowenstam. University of Chicago, Walker Mu-
seum, nos. 45248A and 45248B.

Length of preserved portion of wing, 56 mm.; probable original
length, 60 mm. or slightly more.

NEW FAMILY, UNNAMED

Palaeodictyopteran wings with broad costal area narrowing
distally; Sc, Rl, Rs and MA parallel and running nearly straight
to apex of wing; Rs little divided, forking distally; stem of M curved
backward basally so that Cu and A arise from it; CuP branched;
complete irregular archedictyon.

The single remarkable wing representing this family has several
characteristics normally present in wings of the order Protorthoptera.
Each of these, however, occurs also in one or more genera of Palae-
odictyoptera, though their occurrence together is unique in the order.
Characters that ally the wing to the Palaeodictyoptera are: the
backward curving of MP, Cu, and A; the irregular archedictyon;
the three anal veins; the branched CuP. Each of these characters
may be found in wings of other orders, but their coincidence can
only require an assignment to the Palaeodictyoptera. Since there
is at present only a single specimen displaying this group of charac-
ters, a formal name is not proposed for the family.

Turnbullia, new genus

Description. — Wing medium-sized, about 23^ times as long as
wide, greatest width in distal half. C marginal, with heavy hairs
or spines on its anterior edge in the basal part of the wing. Costal
area broadest in basal part of wing, occupied by irregular archedic-
tyon and oblique veinlets. Sc, R, and M arising independently,
arched at base, running nearly parallel up to Mf, MA then con-
tinuing parallel to Rs. Rf at about mid wing; Mf in distal half of
wing. MA simple, curving only slightly toward posterior margin.
MP sending off branches toward apex of wing. Cu arising near
base from stem of M; CuA simple, arising near origin of Cu; CuP

28 FIELDIANA: GEOLOGY, VOLUME 12

with its posteriorly directed branches occupying half of the posterior
border of the wing. Anal veins arising from stem of M, arching to
posterior margin.

Genotype. — Turnbullia priscUlae, new species.

Remarks. — The genus carries the name of Mr. and Mrs. William
D. Turnbull, the former Assistant Curator of Fossil Mammals,
Chicago Natural History Museum, who have added greatly to the
Museum's fossil collections through preparing and caring for the
specimens, as well as through collecting. The specimen of the
genotype was found by Mrs. Turnbull on one of their collecting
excursions.

Turnbullia priscillae, new species

Figures 8, 9

Description. — Wing, as preserved, 59 mm. long, 25 mm. wide,
probably about 63 mm. long in life. Costal veinlets more regular
in narrow apical portion of costal area than basally. Rs branching
near apex of wing into R2+3 and R4+5; a second forking separates
R4 from R5 very close to the apex. MA arising as a neutral vein
from the compound stem M somewhat distad of Rf ; MP continuing
the direction of the stem and sending off a pectinate series of three
branches toward the posterior slope of the apical margin. CuA
weakly convex. CuP with a pectinate series of four branches
curving toward the posterior margin. lA forked, weakly concave,
arising from the stem of M somewhat basad of the origin of Cu.

Remarks. — Convexity is of little assistance in homologizing the
veins of this species. Rl is strongly convex, as is to be expected, but
the vein designated MA is neutral rather than convex. An alterna-
tive interpretation is that MA is lacking and that the vein following
Rs is a branch of MP; that would bring the wing closer to the
Protorthoptera, to which several other characters seem to draw it.
This relation would be further strengthened by regarding the vein
designated CuPl as CuA, with branches arising from both sides of
its stem. The vein here called lA would then have to be understood
as CuP, a simple concave vein, as in the Protorthoptera, with lA
branching from it. These changes in interpretation, plus the broad
costal area, the straight course of the anterior veins, and the fusion
of other veins onto the base of M, would constitute a strong argument
for placing this wing among the Protorthoptera. However, the stems
here called Cu and lA are probably not far removed from a condition

Fig. 8. Tumbullia priscillae, new species; X 1.6. lA, first anal; C, costa;
CuA, anterior cubitus; CuP, posterior cubitus; M, media; MA, anterior media;
Mf, fork of media; Rl-5, branches of radius; Rf, fork of radius; Sc, subcosta.
Weight of lines indicates relative strength of veins.

Fig. 9. Tumbullia priscillae, new species, holotype; CNHM no. PE149. The
lower half of the concretion, illuminated from below, appears in reversed relief.

29

30 FIELDIANA: GEOLOGY, VOLUME 12

of primitive independence, having been captured by a posterior
migration of the stem of M. The anterior branch of Cu, the only
convex vein in the cubital neighborhood, can logically represent the
normally convex CuA, while the other branches, neutral, must be
the fundamentally concave CuP and its ramifications. The dis-
tinctness of MA and MP is scarcely indicated by the relief of the
veins, but while the branches here labeled MP lie slightly below the
level of the wing membrane and are thus weakly concave the
anterior branch, though actually neutral in relief, lies higher and
may therefore be understood as representing the fundamentally
convex MA.

The very slight degree of branching of Rs is rare; it occurs in
Fabrecia Meunier and Stenoneura Brongniart among the Protor-
thoptera, and in Macroptera Laurentiaux among the Palaeodic-
tyoptera. Laurentiaux (1949b, p. 219) has justly called the retarded
branching of Rs a primitive character.

The major veins and branches retain marks of trichia, indicated
as dots in figure 8. Trichia are present also in the membrane within
the mesh of the archedictyon, having the same size as those on the
veins. Tillyard (1918, pp. 631-634) has shown that macrotrichia
occur only on true veins and veinlets, and not on the wing membrane
or on cross veins. In Turnhullia, the trichial bases on the membrane
are as large as those on the veins, and must certainly be regarded
as macrotrichia. If Tillyard's rule admits no exception,^ Turnhullia
has an aphantoneuric archedictyon represented by the trichial bases,
plus a new secondarily developed complete archedictyon. Such
a hypothesis falls almost of its own weight; the macrotrichia of
Turnhullia must have developed originally on the wing membrane
itself.

The posterior border of the wing is delimited by a fairly strong
marginal vein for which there is no accepted term in the nomen-
clature of veins. It may represent an anal vein that blends apically
with C, or it may be C itself. The wings of modern Mecoptera have
such a marginal vein. The posterior margin of the wing is rippled
by an up-arching of the membrane (and the marginal vein) between
the relatively depressed branches of CuP.

The trivial name is proposed in honor of Mrs. Turnbull, who
found the type specimen and presented it to the Museum.

' Macrotrichia have been reported on cross veins, as by Martynov (1938a,
p. 203); they also occur on cross veins in the modern Planipennia, for example,
in Chrysopa.

RICHARDSON: INSECTS

31

Holotype. — Two halves of a concretion bearing the paired im-
pressions of the wing. CNHM no. PE149. From the strip-mining
area near Coal City (Coordinates d9.0, F4.1 on the map, fig. 3,
this volume, no. 1) ; collected April 26, 1953.

FAMILY UNDETERMINED
UNDETERMINED INSECT NO. 1

Figures 10, 11

A nearly complete body of an insect, lacking wings, but with
parts of four legs. Metathorax and mesothorax about equal in size,
abdominal segments at least nine in number, slightly diminishing in
size posteriorly. Prothorax and head probably present, but charac-
ters obscure. On two of the abdominal segments there are lateral
flaps such as are present on the abdomen of Teneopteron, Stenodidya,
and many other primitive insects. Length, as preserved, 293^ mm.

t ;

Fig. 10. Drawing of
undetermined insect
no. 1.

Fig. 11. Undetermined insect no. 1; X 1.48.
CNHM no. PE3285. The impression of parts of
body and legs of insect among plant fragments.

32

FIELDIANA: GEOLOGY, VOLUME 12

Figured specimen. — CNHM no. PE3285, two halves of a con-
cretion containing abundant macerated plant debris and the paired
impressions of the insect body. Jackson farm, about midway
between Braidwood and Wilmington. (Coordinates h5.5, J7.2, on
the map, fig. 3, this volume, no. 1.) Collected, 1952, by E. S.
Richardson, Jr.

UNDETERMINED INSECT NO. 2

Figures 12, 13

A fragment of a wing, about 20 mm. in length; not sufficiently
complete for description. The fine, close cross veins in subparallel
arrangement give the delicate membrane the appearance of being
wrinkled.

Fig. 12. Drawing of un-
determined insect no. 2; X 3.5.
+ , elevated vein; — , depressed
vein.

Fig. 13. Undetermined insect no. 2; X 1.56. CNHM no. PE146. The
counterpart specimens, illuminated from below, appear in reversed relief.

Figured specimen. — CNHM no. PE146, two halves of a nodule
bearing paired impressions of the wing. Collected by George
Langford in strip mine spoil heap. (Coordinates e7.2, 70.0 on the
map, fig. 3, this volume, no. 1.)

RICHARDSON: INSECTS 33

Order PROTORTHOPTERA

Handlirsch (1937, pp. 63-65) has divided the Protorthoptera
into four suborders, plus an unassigned array of famihes and genera.
Since this division is soon to be superseded by Professor Carpenter's
promised revision, no attempt will be made here to place the follow-
ing species in suborders or to break up the unwieldy order Protor-
thoptera. The first two genera that follow are included in the
family Cacurgidae, and should therefore go in Handlirsch's suborder
Cacurgoidea, though his definition of the suborder is not entirely
appropriate.

Family Cacurgidae Handlirsch 1911, emend. Pruvost 1930

Discussions of the family are to be found in Laurentiaux (1949a,
p. 55) and works there cited. In this family, Handlirsch included
the following genera in 1922: Cacurgus Handlirsch 1911 (Mazon
Creek); Spilomastax Handlirsch 1911 (Mazon Creek); Palaeomastax
Handlirsch 1904 (Westphalian of Belgium) ; Archaeologus Handlirsch
1906 (Mazon Creek); Archimastax Handlirsch 1906 (Pottsville of
Arkansas); Cacurgellus Pruvost 1919 (Westphalian of France). He
properly excluded Oryctomastax Pruvost 1919 and Archaeacridites
Meunier (included by Pruvost in 1919). To the above genera,
Pruvost added (1930) Omalia Van Beneden and Coemans 1867,*
including as synonyms Palaeomastax Handlirsch and Coselia Bolton
1922 (Westphalian, Belgium and England). Carpenter (1943b)
added Heterologus Carpenter 1943 (Braidwood strip mines). It is
now proposed to add two more genera, Nacekomia and Anthrakoris.

Nacekomia, new genus

Description. — Cacurgids with narrow forewing with pectinately
many-branched MP; simple Rl joined to C in distal third of wing
by forking veinlets; Rs twice dichotomously divided; lower branch
(CuPl) of cubito-medial Y-vein very long, leaving a broad space
between CuPl and CuP2; cross veins in medial field very strong;
four strong anal veins.

Diagnosis. — Nacekomia, like the other genera of the family,
differs from Cacurgus in its strong and relatively straight and distant
cross veins, with only a tendency to develop the polygonal network

' Scudder (1885, p. 331), having seen the original figure of Omalia macroptera,
remarked that "its curious venation is plainly impossible."

34 FIELDIANA: GEOLOGY, VOLUME 12

seen in the latter genus. It differs from Cacurgus also in its simple
R, its classically dichotomous Rs, and its greatly branched pectinate
MP. In a comparison of the two genera, the media of Nacekomia
may be said to take the place of the cubitus of Cacurgus in the
structural scheme of the wing, as does the radial sector in Heter-
ologus. Cacurgus and Nacekomia are similar in possessing a large
area in the cubital field unsupported by a major vein, due in part
to the far basal position of Cuf with respect to Mf. Spilomastax,
Archimastax, and Archaeologus are insufficiently known, but are
similar to Nacekomia in the relation of Cu to M. Cacurgellus seems
to have a simple, strongly concave CuP (which Pruvost regarded
as an anal) . Heterologus and Nacekomia both have R2 fused distally
with Rl, and have also a nearly straight anterior margin, but Heter-
ologus lacks the area of confused venation in the cubital field.
Omalia, similar in this respect, differs greatly in the branching and
relations of Rl and M. Nygmata (wing spots) are absent. Nace-
komia is the only genus of the family known to have a strongly
sclerotized area at the base of the wing. The forewing of Archaeo-
logus apparently had proportions similar to those of Nacekomia, but
had the cubital fork close to the medial fork, as in other cacurgids.

Remarks. — A "barbarous name," as contemplated in Recom-
mendation j of the International Rules of Zoological Nomenclature,
being a Latinization of the Russian word for "insect."

Genotype. — Nacekomia rossae, new species.

Nacekomia rossae, new species
Figures 14, 15

Description. — Forewing more than three times as long as wide,
greatest width in distal half of wing. Humeral area strongly convex,
sclerotized, with convex margin; anterior margin of wing broadly
emarginate in proximal part, broadly convex in distal part. C mar-
ginal, strong. Sc terminating on C at about one-third the distance
from the apex. Rl and CuA the only strongly convex veins in the
wing. Cross veins strong, numerous, nearly perpendicular to
branches of longitudinal veins.

Holotype. — CNHM no. PE791, one half of a concretion bearing
the impression of the under surface of the wing. Length of wing,
as preserved, 43 mm.; width in distal half, 12 mm. The apex of the
wing evidently continued for at least a millimeter beyond the edge
of the concretion, which had been slightly reduced in size by weath-

RICHARDSON: INSECTS

35

ering before splitting open naturally. Collected in 1948 by Miss
Lillian Ross, Associate in Insects, Chicago Natural History Museum,
and presented by her to the Museum. (From the strip-mine area.)
Remarks. — The humeral area, devoid of costal veinlets, is smooth
and strongly convex, evidently sclerotized. An elevated or depressed,
smoothly rounded area, indicated by a pattern of dots in the drawing

CuPI • CuA +

Fig. 14. Nacekomia rossae, new species; X 2.5. A, anal; C, costa; CuA,
anterior cubitus; CuPl, CuP2, first and second forks of posterior cubitus; MP,
posterior media; Rl-2, branches of radius; Rs, radial sector; Sc, subcosta. +,
elevated vein; — , depressed vein. Weight of lines indicates relative strength of
veins. Dotted area, sclerotized.

Fig. 15. Nacekomia rossae, new
species, holotype; X 1.13. CNHM
no. PE791.

(fig. 14), continues this basal homy portion to the edge of the anal
field and also distally as a projection covering the bases of the
pre-anal, post-costal veins. The rest of the wing membrane was
very thin, the texture of the specimen showing a minute wrinkling.
The strong cross veins of the medial field bear transverse striae,
about 50 per millimeter. The under surface of Sc is marked by
small irregular diagonal impressed lines possibly representing pros-
trate trichia.

The homologies of the veins have been determined with the help
of the argument used by Carpenter in his valuable discussion of the
veins of Caloneura dawsoni (Carpenter, 1943a, p. 69) ; since the only
vein in medial position is concave, it must be MP; MA, as in Calo-
neura and very many other orthopteroids, must be lacking. The

36 FIELDIANA: GEOLOGY, VOLUME 12

horny base of the wing is convex over the stem of M and Cu, and
may represent the missing MA. However, being a speciaHzed
structure, it probably does not represent a primitive topography.
CuP branches into two concave stems very near the base, within
the heavily sclerotized area, CuPl then running parallel to the
anterior longitudinal veins to its junction with CuA; CuP2 parallels
the course of the anal veins, and the wide gap between these two
branches of CuP is occupied by irregular veins of indefinite homology.
The unusual strength of the cross veins in the medial field makes
a comparison with the caloneurodeans tempting, but the structure
of Cu precludes placing Nacekomia in that order. The presence of
only two prominent convex veins, Rl and CuA, is common in most
insect orders (Imms, 1934, p. 39).

Anthrakoris, new genus

Description. — Anterior margin of forewing rounded, posterior
margin nearly straight. Costal area broad, occupied by branching
veinlets with cross veins. Sc terminating about one-fourth of the
wing length from the apex. Rs arising slightly basad of mid wing,
with few branches. MA absent; MP arising close to or coalesced
with R at base, with many branches. CuA and CuP coalesced at
base. CuA forked, CuP simple, nearly straight. Two unbranched
anal veins. Cross veins not plentiful; mesh-like archedictyon
present in basal half.

Genotype. — Anthrakoris aetherius, new species.

Remarks. — The shape of the wing in this genus is most like that
of Omalia, among the cacurgids, a genus from which it differs in
lacking the fusion of Rs with MP, and in the lesser development
of cubital branches. The termination of Sc on Rl is a character
shared with Heterologus. The venation of Anthrakoris is very much
like that of the palaeodictyopteran Aenigmatodes Handlirsch 1906.
The differences, however, are significant, and the resemblance is due
simply to convergence.

Anthrakoris aetherius, new species
Figures 16, 17

Description. — Forewing: Length, as preserved, 36 mm., entire
length probably not over 40 mm.; width, 13 mm. Anterior margin
strongly convex, greatest curvature near base; posterior margin only
slightly curved, very slightly emarginate at the end of CuP; greatest

RICHARDSON: INSECTS

37

Rs-

MP-

FiG. 16. Anthrakoris aetherius, new species. lA, first anal; 2A, second anal;
C, costa; CuA, anterior cubitus; CuP, posterior cubitus; MP, posterior media;
Rl, principal branch of radius; Rf, fork of radius; Rs, radial sector; Sc, subcosta;
+, elevated vein; — , depressed vein. Weight of lines indicates relative strength
of veins; archedictyon indicated only in part.

Fig. 17. An</iraA;omac</ienMS, new species, holotype; X 1.7. CNHM no. PE145.

width at about midwing. Sc close and parallel to Rl, both doubly
sigmoidally curved; Rs emitted shortly basad of midwing at most
posterior part of curve of R. Branches of Rs and MP occupy
a subtriangular area whose base is the apical margin of the wing and
whose sides are defined by the prominent convex veins Rl and CuA,
forming a very characteristic pattern. Besides these two convex
veins, the most conspicuous vein in the wing is the deeply set

38 FIELDIANA: GEOLOGY, VOLUME 12

concave CuP, occupying a plical furrow. Details of the venation
are apparent in the figures.

Holotype. — Two halves of a concretion bearing the impressions
of the dorsal and ventral surfaces of the wing. Received at the
Museum as a gift of the University of Chicago, 1951, included in the
fossil plant collection of the late Professor A. C. No^. The lithology
of the specimen is of the Mazon Creek type, rather than the strip
mine type. CNHM no. PE145.

Family Blattinopsidae Bolton 1925

Handlirsch (1937, p. 64) attempted to show that this family
should properly be termed Oryctoblattinidae, the name under which
he had included it in his Fossilium Catalogus (1922). This view,
however, is apparently based on a misunderstanding of the species
proposed by Scudder as genotype of Oryctoblattina, and Bolton's
name should prevail.

Glaphyrokoris, new genus

Diagnosis. — Similar to Glaphyrophlehia Handlirsch 1906, but
with fairly regular cross veins in place of a meshwork between the
longitudinal veins, and with MP arising from the stem of R rather
than independently.

Genotype. — Glaphyrokoris mirandus, new species.

Glaphyrokoris mirandus, new species

Figures 18, 19

Description. — FoREWlNG: Anterior margin strongly arched,
posterior margin nearly straight. C marginal, costal space broad,
occupied by many slanting veinlets arising from Sc. Sc terminating
on C at about midwing. R simple, distant from Sc in basal half of
wing; close to anterior margin beyond end of Sc, terminating on
anterior margin. Rs arising in basal half of wing, distant from Rl
in apical half, sending forth a pectinate series of four posteriorly
directed branches. MA lacking; MP arising near base from stem of
R, forking twice at about midwing. M5 present as a short convex
vein joining MP to Cu near origin of MP. CuA strongly convex,
arising independently, emitting many branches curved toward the
apex of the wing. CuP simple, in deep plical furrow; posterior
margin of wing not inflected at end of furrow. lA strongly convex,

RICHARDSON: INSECTS

39

simple; 2A with three short curved branches concave toward the
apex of the wing; 3 A with at least one similarly curved branch.

Length, as preserved, 153/^ mm.; original length probably 19 or
20 mm. Greatest width, 73^ mm. This wing is about twice the size
of the forewing of Glaphyrophlebia pusilla Handlirsch, which it re-
sembles in general aspect.

2A lA CuP

CuA

Fig. 18. Glaphyrokoris mirandus, new species. 1-3A, anals; CuA, anterior
cubitus; CuP, posterior cubitus; MP, posterior media; Rl, principal branch of
radius; Rs, radial sector; Sc, subcosta.

Fig. 19. Glaphyrokoris mirandus, new species, holotype; X 1.6.
no. PE977.

CNHM

Holotype. — CNHM no. PE977, two halves of a small concretion,
bearing impressions of the wing. Collected in 1949 by George
Langford. (Coordinates e7.2, 70.0 on the map, fig. 3, this volume,
no. 1.)

40 FIELDIANA: GEOLOGY, VOLUME 12

FAMILY UNDETERMINED

UNDETERMINED INSECT NO. 3
Figures 20, 21

A large imperfectly preserved protorthopteran insect, known
from only one half of a concretion that had opened by weathering.
Prolonged search in the vicinity failed to reveal the missing counter-
part, which should have borne the impression of the entire wings.
Although the specimen may be referred to the rather indefinite order
Protorthoptera, it is not assigned to a family or genus. It may very
likely fall within Handlirsch's suborder Geraroidea.

Body, consisting of head, the three thoracic segments, and five
of the abdominal segments, 54 mm. long, as preserved. Forewings
at least 50 mm. long; wing-spread at least 108 mm. Head large,
medially constricted, widest posteriorly, where it embraces the pro-
thorax, flaring anteriorly and there terminated as though cut ver-
tically; a raised border on the front margin, rising to a blunt spine
on the center. In front of the head is a flattened impression of
indefinite triangular shape, evidently representing some part of the
insect, such as lies in front of the body of Teneopteron mirabile.
Prothorax slightly longer than wide, expanding anteriorly into two
lobes, and terminating at the back in a marginal ridge with a prom-
inent blunt central dorsal protuberance. Metathorax subtrap-
ezoidal, tapering to the rear, its greatest width greater than its
length; a low central dorsal protuberance but no elevated border on
the rear margin. Abdomen tapering to the rear. Two forelegs with
sturdy femora.

Costa of forewing marginal; veinlets in costal area straight,
slightly oblique, joined by cross veins in basal part of wing, straight
or curved, oblique, forked in apical portion, as in Anegertus. Sc
long, about parallel to C, and evidently terminating on C at some
point distad of midwing. Rl simple; Rs arising at about midwing
and not forked within the portion observable. Cross veins straight,
oblique or perpendicular to longitudinal veins. Distal margin
unknown; anterior margin only slightly bowed. Hindwings repre-
sented by only a very small fragment.

A complete specimen in the collection of Mr. Peter Enrietta, in
Coal City, Illinois, may be conspecific with this insect.

Figured specimen.— CNRM no. PE2919. Collected April 30,
1952, by E. S. Richardson, Jr. (Coordinates e7.2, 72.0 on the map,
fig. 3, this volume, no. 1.)

RICHARDSON: INSECTS

41

Fig. 20. Drawing of undetermined insect no. 3, partially restored; X 1.3.

Fig. 21. Undetermined insect no. 3; X 0.86. CNHM no. PE2919.

Order CALONEURODEA

For a discussion of the order, here reported in its earliest known
occurrence, see remarks below.

Genopterygidae, new family

This family is proposed for two monotypic genera, Genopteryx
Scudder 1885 and Rossites, new genus. Genopteryx clearly has the
two characters regarded by Carpenter (1943a) as prescribing in-
clusion in the order Caloneurodea : the heavy cross veins and the
close straight parallel CuA and CuP. Rossites has delicate cross
veins and its CuA deviates from strict parallelism with CuP, yet
the venation is nearly identical with that of Genopteryx constricta.
Except for the importance to be assumed in the later development

42 FIELDIANA: GEOLOGY, VOLUME 12

of the Caloneurodea by the heavy cross veins, these two insects
might quite properly be included in one genus.

Family characters are: Rs much divided; MP and CuA arising
from Rs; CuP arising from lA; 2A arising from lA.

The two genera of the Genopterygidae are the oldest members
of the order known at present, and in their similarities and differ-
ences point to the manner of origin of an order in its first breaking
away from an ancestral stock. In the variation within the group
of the Protorthoptera-Protoblattoidea, a combination of characters
was attained much like those of Rossites. Further variation added,
in the case of Genopteryx, a single new character, the heavy cross
veins, at the same time affirming and emphasizing the parallelism
of CuA and CuP. At this point, the difference between Genopteryx
and Rossites could well be considered specific rather than generic.
But some selective advantage appears to have been bestowed upon
Genopteryx and its descendants along with the adoption of the
heavy cross veins. Rossites passed out of the picture at the end of
one of the innumerable culs-de-sac of evolution, while the seed of
Genopteryx spread in various directions, forming a well-defined
group properly to be regarded as an order. Within the limits imposed
by the two ordinal characters, which remained remarkably constant
through the Pennsylvanian and the Permian, the number of possible
variations is not large, though greater than was achieved in the
presently known genera of the order. The general trends may be
seen in comparing the wing patterns of the caloneurodean genera
now recognized: the termination of Sc on C migrates basad, as in
the Homoptera; the number of branches of Rs is reduced; the number
of anal veins is reduced and the wings become subpetiolate ; Rf
moves apicad; the origins of CuA and MP vary from one stem to
another, probably in adjustment to the narrowing of the wing and
to the change in position of Rf. These changes proceed at different
rates in several different lines, and on the basis of the few known
genera, a number of hypothetical phylogenetic trees may be drawn
to express this. Since, however, any one of those trees seems as
valid as any other, none is represented here.

Scudder (1885) included Genopteryx in his "Neuropteroid
Palaeodictyoptera," family Homothetidae, from which Handlirsch
removed it to his family Geraridae, in the Protorthoptera. Mar-
tynov (1938b, p. 99) transferred the Geraridae to the order Para-
plecoptera, which Carpenter (1943a, p. 75) regards as a doubtful
unit. Since the disposition of these entities will no doubt be in-

RICHARDSON: INSECTS 43

eluded in Professor Carpenter's revision of the Protorthoptera, they
will not be further discussed here.

Genus Genopteryx Scudder 1885

Genopteryx Scudder, 1885, Mem. Boston Soc. Nat. Hist., 3, p. 327.

Scudder's definition of the genus is repeated here, the names
of the veins according to the current nomenclature being introduced

Fig. 22. Genopteryx constricta Scudder. lA, first anal; C, costa; CuA,
anterior cubitus; CuP, posterior cubitus; MP, posterior media; Rl, principal
branch of radius; Rs, radial sector; Sc, subcosta. Drawing by Laura Sparks from
holotype, USNM no. 38148.

in brackets, together with a mention of the convexity, as determined
by examination of the type specimen of Genopteryx constricta
Scudder (fig. 22), seen through the courtesy of Dr. G. Arthur Cooper,
United States National Museum.

Wings obovate, with more or less arched costa [C, convex] and somewhat
produced apex; mediastinal vein [Sc, concave] of variable length, the scapular
[Rl, convex] extending to or nearly to the tip, connected to the veins on either side
of it by transverse or oblique cross veins; externomedian vein [M; actually Rs,
concave, plus MP, concave, plus CuA, convex] very important, commencing to
branch considerably before the middle of the wing and by several longitudinally
oblique mostly forked veins, closely connected by feebler cross veins, feeding the
apex of the wing; internomedian vein [Cu; actually CuP, concave, and the anals]
also important with several similar veins [the anals?], the outermost of which
[CuP] runs in close proximity to the basal externomedian branch [CuA] from
its very origin, so that at first sight both externomedian and internomedian
branches appear to spring from a common vein.

The genus apparently includes the single species Genopteryx
constricta Scudder 1885. Scudder originally included also Gryllacris
lithracantha Goldenberg, but Handlirsch in 1906 properly removed
it to another genus.

44 FIELDIANA: GEOLOGY, VOLUME 12

Rossites, new genus

This genus is proposed for the reception of a single species
closely related to Genopteryx constricta but differing in lacking
strongly developed cross veins.

The genus is named in honor of Dr. Herbert H. Ross, of the
Illinois Natural History Survey, to whom I am indebted for several
helpful conversations; and his son, Mr. Charles A. Ross, who col-
lected a specimen of the genotype.

Genotype. — Rossites inopinus, new species.

Rossites inopinus, new species

Figures 23-25

Description. — Wing: Costal area of average width, occupied by
veinlets increasing in obliquity toward the apex of the wing. Sc
terminating on C in apical half of wing. Base of Rl very close to
base of Sc, but Rl diverging from Sc near base so that costal and
subcostal areas are of about the same width. Rs arising very near
base of wing; Rs with 5 or 6 pectinately arranged oblique branches
occupying the apical portion of the wing. MP arising from Rs,
forked. CuA, strongly convex, arising from stem of Rs, continuing
the curvature of that stem until it nearly touches CuP, at which
point it is angulated and thence continues more distant from CuP,
about straight, to the posterior margin; simple or forked. CuP
arising independently of CuA, strongly concave, nearly straight,
simple or forked. lA arising from stem of CuP; 2A (or a branch of
lA?) arising near base from the same stem.

Holotype. — Two halves of a concretion containing impressions
of the right wing. Collected in 1949 by George Langford. The
apical portion of the wing is missing; the remaining portion is 28
mm. in length. CNHM no. PE981. (Coordinates e7.2, 70.0 on map,
fig. 3, this volume, no. 1.)

Paratype. — One half of a concretion bearing the impression of
the dorsal surface of a right wing. Collected by Charles A. Ross
and presented by him to the Museum. CNHM no. PE3304.
(Coordinates approximately e7.2, Jl.O on map, fig. 3, this volume,
no. 1.)

The paratype differs from the holotype in being about 1 3^ times
as large, in lacking a faint fork on lA, and in that CuA is forked and
CuP not.

RH-

0 1

10
13 mm.

Fig. 23. Rossites inopinus, new species; A drawn from holotype, B from
paratype. lA, first anal; C, costa; CuA, anterior cubitus; CuP, posterior cubitus;
MP, posterior media; Rl, principal branch of radius; Rs, radial sector; Sc, sub-
costa; +, elevated vein; — , depressed vein. The dot-and-dash line in A indicates
a break in the wing membrane, whereby the anal area may not be in correct
relation to the rest of the wing.

V

Fig. 24. Rossites inopinus, new species, holotype; X 1. CNHM no. PE981.
Associated markings in concretion are indeterminate plant debris.

Fig. 25. Rossites inopinus, new
species, paratype; X 1.1. CNHM
no. PE3304. An impression of upper
surface of wing, veins appearing in
reverse relief.

45

46 FIELDIANA: GEOLOGY, VOLUME 12

ORDER UNCERTAIN

The following new family is based on a species that Carpenter
(1943b, p. 17) called "probably the most remarkable insect which
has been found in the Mazon Creek nodules." In ordinal position
it probably lies closest to the Blattaria in wing venation, but,
lacking the characteristic modification of the blattarian prothorax,
it cannot be included within that order. It bears a superficial
resemblance to Idelia M. D. Zalessky, from the Russian Permian
(order Protorthoptera) , but is more specialized in several respects;
hence it cannot lie in that lineage. It can not be considered ancestral
to the Permian order Protelytroptera, another order in which the
fore wings are heavily sclerotized, as its venation is already too
highly specialized. In shape, relative size, and generalities of
venation, the wings resemble those of the modern Tettigoniidae
(Orthoptera) .

Teneopteridae, new family

Medium-sized neopterous insects with slender tegminous fore-
wings, the principal longitudinal veins grouped in a band on or near
the midline, many veinlets occupying the costal area, the anal area
set off by a plical furrow; legs robust, homonomous; abdomen no
longer than thorax.

Genus Teneopteron Carpenter 1943

Teneopteron Carpenter, 1943, Illinois State Mus., Sci. Pap., 3, pt. 1, p. 17.

The genus was defined only by comparison with Megagnathites
(Bolton) Handlirsch, to which, however, it is not related. Characters
of generic importance are probably the following, combined with the
family characters suggested above: Prothorax with elevated rachis
in anterior half, rachis expanding anteriorly, divided by median
groove; a pair of low tubercles on posterior half of prothorax.
Mesothorax shorter than other thoracic segments. Forewings more
than three times as long as wide, sclerotized, nearly flat antero-
posteriorly, with convex humeral area; principal longitudinal veins
arising at base in two deep grooves, the posterior groove arching
backward as a plical furrow; Sc terminating on C in distal third
of wing.

Genotype. — Teneopteron mirahile Carpenter 1943 (by original
designation) .

RICHARDSON: INSECTS 47

Teneopteron mirabile Carpenter 1943
Figures 26-30

Teneopteron mirabile Carpenter, 1943, Illinois State Mus., Sci. Pap., 3, pt. 1,
pp. 17-20.

This species was founded on a single specimen, no. 14887 in the
Ilhnois State Museum. In describing it, Carpenter, misled by the
extraordinary venation, thought that the forewings had been over-
turned, and accordingly interchanged anterior and posterior in his
description. The second specimen, no. PE967 in Chicago Natural
History Museum, is somewhat better preserved than the type, and
clearly shows the position of the wings. Thus, the species may now
be more fully described. The following description and discussion
are based on the two specimens, which are interpreted as conspecific
in spite of certain apparent differences. Even though the specimens
should in the future be considered distinct species, they are so
closely related that the orientation of the wings of the original
specimen can no longer be in doubt.

Forewings. — The forewings are thick and sclerotized, nearly flat
transversely, with a convex humeral area. Longitudinally, they are
bowed upward. The anal area, probably of thinner membrane,
is bent at the plical furrow out of line with the rest of the wing
surface. Apparently it is the same sort of anal area as appears in
the forewings of modern Tettigoniidae, and the manner of folding
the wings was probably similar, with the small anal area folded
down over the mesothorax.

The veins of the forewing arise in two furrows at about the
middle of the base. Homologies of the veins, indicated in the ac-
companying drawings (fig. 26), are tentative, as the veins stand
in very low relief and many can be seen only with special lighting.
What veins may be expected in the order are, of course, not known.
The humeral area is clear of veinlets, and is succeeded by the costal
area, occupied by many curved veinlets arising from Sc. The
anterior edge of the wing just beyond the humeral area is emarginate
and minutely serrated. Sc apparently curves into the costal margin
at about two-thirds the length of the wing. Beyond its termination,
veinlets from Rl continue the pattern of branches reaching the
anterior margin. The longitudinal vein denoted as Rs sends off
branches to the posterior margin in its distal half. As far back
toward the base of the wing as it can be separately distinguished,
it is independent of R. The veins already mentioned arise in the
anterior of the two grooves at the base of the wing; two succeeding

Fig. 26. Teneopteron mirabile Carpenter. A, drawn from holotype, slightly
restored. B, drawn from plesiotype, slightly restored. A, anal; CuA, anterior
cubitus; M, media; R, radius; Rs, radial sector; Sc, subcosta.

48

RICHARDSON: INSECTS

49

Fig. 27. Teneopieron mirabile Carpenter, plesiotype; X 1. CNHM no.
PE967, consisting of counterpart impressions.

Fig. 28. Teneopteron mirabile Car-
penter, holotype; X 1. Illinois State
Museum no. 14887.

veins, designated M and CuA, arise from the other groove, which
corresponds to the plical furrow, containing CuP, of the roaches.
Within the anal area, behind the pHcal furrow, is a series of extremely
fine lines radiating from a point posterior to the basal origin of the
furrow and directed to the posterior margin of the wing basad of
the outer end of the furrow. This is the character of the anal veins
in the most primitive Blattaria, as emphasized by Tillyard (1919,
p. 362; 1937b, pp. 177-178), though in the present specimen (CNHM
PE967) they are unusually straight and unusually weak. The
straightness is probably, then, a primitive character.

Hindwings. — The distal tips of the two hindwings are visible
in the CNHM specimen. From them, little of the character of the
wing can be made out. The veins all stand in relief on the ventral

50

FIELDIANA: GEOLOGY, VOLUME 12

Fig. 29. Teneopteron mirabile Carpenter; X 6.1.
CNHM no. PE967, showing venation of hindwings.

Portion of plesiotype,

surface of the wing and are impressed on the dorsal surface. The
membrane in this distal portion of the wing is flat, without concave
and convex veins, and with no cross veins. The hindwings are
somewhat longer, from base to tip, than the fore wings, and, since
they originate farther back on the body, it is evident that they must
have projected from beneath the ends of the forewings if they were
not folded when in repose. The Coleoptera and the Protelytroptera
protect the long hindwings by folding the apical ends upward; the
Tettigoniidae and perhaps Teneopteron permit the distal end of
the hind wing to project beyond the end of the tegmen. The hind-
wing has a broad costal area occupied by slanting veinlets, in the
orthopteroid fashion; the tip of the wing is broadly rounded.

Thorax. — Three segments of about equal size, the mesothorax
being the shortest. Crushing and distortion have been most trouble-

RICHARDSON: INSECTS

61

Fig. 30. Teneopteron mirabile Carpenter; X 3.85. Detail of thoracic region
and wings of plesiotype; CNHM no. PE967.

some in these segments, so that they cannot be described in detail.
The prothorax has the characters described for the genus.

Abdomen. — The abdominal segments are very clearly preserved
in both specimens, with the exception, in both, of the posteriormost
portion. In the holotype, the one or two final segments form
a smoothly rounded termination of the body, with no cerci or other
appendage. Carpenter drew and described this as two clearly dis-
tinct segments, which it must surely be. Counting it as two, the
number of abdominal segments is nine. In the plesiotype there is
apparently a central boss on the seventh segment (it may be merely
a small rosette of marcasite), followed by a medial groove in the last
two segments.

The two specimens are drawn as they appear under the micro-
scope (fig. 26), the apparent differences probably being due as
much to preservation as to natural variation between the specimens.
On both, the abdomen gradually widens posteriorly to the sixth
segment, the last three being narrower. The segments are about
fiat dorsally, with angulated edges and steep pleural slopes. The
edges of the terga in the holotype are set off by low ridges. In both
specimens, a low ridge traverses the first seven segments longitudi-

52 FIELDIANA: GEOLOGY, VOLUME 12

nally; this is single in the plesiotype but double in the holotype.
In the holotype it continues through the three terminal segments,
but in the other it ends against the boss on the seventh segment.
Carpenter regarded the portion of the tergum between the paired
ridges as a separated median plate. The difference in appearance
between the two specimens almost certainly does not reflect an
anatomical difference, as the appearance of the under side of the
abdomen may vary extremely in individual specimens of dried
tettigoniids, both of these aspects and many others being among
the possible configurations of wrinkles in this region.

Paired lateral abdominal processes, which Carpenter interpreted
as gills signifying an aquatic larval stage, are visible on all abdom-
inal segments of the holotype but are not visible on the other spe-
cimen; their lack is probably due to imperfect preservation.

Legs. — Parts of the femora of all six legs are visible on the
CNHM specimen and four on the other. The trochanter and coxa
are present on the two hind legs of the plesiotype, and on one hind
leg of the other, but are so crushed or wrinkled that it is not possible
to distinguish their details. A nearly complete tibia is articulated
to the femur of a hind leg of the type specimen. Femora robust,
those of the forelegs probably less so than the others; tibia of hind
leg slender, shorter than femur.

Measurements. — Holotype: body, front of prothorax to end of
abdomen, 22 mm.; thorax, llj^ mm.; abdomen, 10%! mm.; forewing
(restored), 16 mm. Plesiotype: body, front of prothorax to end of
abdomen, 27 mm.; thorax, 14 mm.; abdomen, 13 mm.; forewing,
193/^ mm.

Locality. — The two specimens were found about 50 yards apart,
on opposite sides of the section line road, 3 miles north of Braidwood.
(Coordinates e7.0, H6.3 [holotype] and eY.O, H9.8 [plesiotype], on
the map, fig. 3, this volume, no. 1.) Both were collected by IVfr.
George Langford.

Plesiotype. — CNHIVE no. PE967, two halves of a concretion
bearing paired impressions of the insect.

NYMPH

Figure 31

This specimen pertains to an undetermined order. It has much
the appearance of a palaeodictyopteran.

RICHARDSON: INSECTS

53

Fig. 31. Nymph; X 1.36. CNHM no. PE973. Associated material includes
plant fragments and a large crustacean claw.

The mesothorax, metathorax, and parts of five abdominal seg-
ments of a rather large immature insect are preserved in a con-
cretion, associated with two legs of a large, undetermined crus-
tacean. The insect specimen undoubtedly represents a late instar
in the larval development of a winged hemimetabolous form. The
mesothorax is about square, 73^ mm. on an edge; the metathorax,
with the same width, is 63^ mm. long; the abdominal segments are
about 2 mm. long. A median groove in each of the thoracic segments
may have been deepened by the pressure that evidently flattened
the specimen, and the width of the body exaggerated, but the groove
is a character of the species, since the abdominal segments, subjected
to the same pressure, have no such groove.

The four large and heavy wing pads are immovably fixed to the
thoracic segments, though the position of the future axillary sclerites
is indicated by a modeling of the surface of the integument at the
junction of the body with the wing pads. Venation is not developed
but is foreshadowed by a pair of deep but indistinct furrows at
the base of each pad, probably representing Sc and CuP of the
adult wing.

The wing pads are laid back along the edge of the abdomen
at a low angle to the body. They are initially directed outward

54 FIELDIANA: GEOLOGY, VOLUME 12

beyond the edge of the thorax before turning along the abdomen,
and thus do not cover the body. By the time of the latest larval
instars, the wing pads of modern insects have developed venation.
The position of the larval wings in this fossil nearly resembles that
of the super-ordinal group Opisthoptera of Lemche (1940, pp.
147-154, fig. 5, 6), distinct from the developmental stages of pla-
giopterans such as the Palaeodictyoptera (op. cit., fig. 5, a).

The specimen resembles in a general way the supposed palaeo-
dictyopterous larvae discussed by Carpenter (1948), who removed
the five previously described specimens from that order for various
reasons. The lack of venation and the fragmentary state of the
specimen at hand make it impossible to assign this larva to a known
order and fruitless to give it a specific name.

Figured specimen. — CNHM no. PE973, two halves of a con-
cretion bearing matching impressions of the nymph. Collected by
Mrs. Robert H. Whitfield and presented by her to the Museum.
(Coordinates e7.0, /i/9.8, in the map, fig. 3, this volume, no. 1.)

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