The wing venation and systematics of Lower Permian Diaphanopterodea

from the Ural Mountains, Russia (Insecta: Paleoptera)

JARMI
LA KUKALOVA-PECK
Ont., Canada KIS 5B6
Department of Earth Sciences, Carleron University, Orravt~~,
AND

N I N AD. SINICHENKOVA
Paleontological Institute, Academy of Sciences, Profsoyuznayu 123, 11 7868 Moscow V-321, Russia
Received November 26, 1990
Accepted August 16, 1991
Can. J. Zool. Downloaded from www.nrcresearchpress.com by Capital Normal University on 03/17/19

KUKALOVL-PECK, J., and SINICHENKOVA, N. D. 1992. The wing venation and systematics of Lower Permian Diaphanop-
terodea from the Ural Mountains. Russia (Insecta: Paleoptera). Can. J. Zool. 70: 229 -235.
Extinct Diaphanopterodea are described from the Lower Permian of Tshekarda, Urals, Russia: Parelmoidae: Uralia n.gen.
(Uralia maculata n.sp. and Uralia sharovi n.sp. ); Paruraliidae n. fam. : Paruralia n.gen. (Paruralia rohdendorfi n. sp. and
Paruralia carpenten n.sp.). Of all Paleozoic insects, the exceedingly well-preserved specimens of U. maculara have contrib-
uted the most important clues to the pterygote ground plans of the head, mouthparts, leg segmentation, thoracic and abdominal
pleura, and genitalia.

KUKALOVL-PECK, J., et SINICHENKOVA, N. D. 1992. The wing venation and systematics of Lower Permian Diaphanop-
terodea from the Ural Mountains, Russia (Insecta: Paleoptera). Can. J. Zool. 70 : 229-235.
On trouvera ici la description d'espttces fossiles de Diaphanopterodea du Permien inferieur trouvees a Tshekarda, Oural,
Russie : parmi les Parelmoidae, deux espttces d'Uralia n.gen., Uralia maculata n.sp. et Uralia sharovi n.sp., et parmi les
Paruraliidae n. fam., deux espttces de Paruralia n.gen., Paruralia rohdendorfi n. sp. et Paruralia carpenten n. sp. Parmi tous
les insectes du Paleozoi'que, U. maculata est l'espkce qui a fourni les indications les plus importantes au sujet de la morpho-
logie de base de la tete, des pikes buccales, de la segmentation des pattes, des pleures thoraciques et abdominales ainsi que
For personal use only.

des genitalia des pterygotes.

[Traduit par la redaction]

Introduction Sinichenkova 1987). The shores were covered by forests of

The Diaphanopterodea is an extinct order of Paleozoic
Paleoptera. It shares with the Paleodictyoptera, Megasecop-
tera, and Permothemistida a specialised sucking rostrum with
a sliding mandibular articulation. Diaphanopterodea have an
exceedingly primitive mechanism of wing folding, presumably
close to that of the ancestral Pterygota (Kukalova-Peck 1983,
1985, 199 1). This paper discusses new taxa from the Lower
Permian (Kungurian) of Tshekarda, near Perm, Middle Ural
Mountains, Russia.
The richest and best preserved collection of members of this
order is a total of 60 specimens of Diaphanopterodea from
Tshekarda. The present study is concerned only with 23 speci-
mens belonging to two families of relatively large Permian
Diaphanopterodea, the Parelmoidae and Paruraliidae; the
remaining specimens belong to the family Asthenohymenidae
and will be described in a separate paper.
The name of Uralia maculata (Parelmoidae) described here,
was first mentioned by Sharov (1973) as a nomen nudum in
a preliminary report, but the intended description never fol-
lowed because of Sharov's sudden death in 1973. Subse-
quently, a short reference to Uralia was made by Rasnitsyn
(1980, with a reconstruction of Uralia sp. by A. G.
Ponomarenko, Fig. 20) and also by Kukalova-Peck (1983,
Fig. 1; 1985, Figs. 33A, 33B).
Tshekarda was discovered as a locality for collecting by
G. T. Mauer in 1930, and subsequently, G. M. Zalessky and
especially A. G. Sharov, who also assembled the specimens
described below, collected there (Martynov 1938; Sharov
1973). The entire sample consisted of more than 5000 speci-
mens of fossil insects. The insects are preserved in the calcare-
ous muds of a brackish, river-fed lagoon (Kalugina 1980;
early gymnosperms (Cordaitales), pteridosperms, and ferns.
The climate was probably warm temperate and relatively
moist. The occurrence of many complete insect bodies indi-
cates rapid burial and the presence of few or no predators or
scavengers.
The Tshekarda locality is renowned for excellent, often
three-dimensional preservation of complete insects, showing
minute morphological details such as hairs, ocelli, articula-
tion, mouthparts, exites, and endites. The dark coloration was
not lost during fossilisation, and the color markings on wings
and sclerotised or dark structures stand out distinctly. On occa-
sion, fossilisation created an "X-ray image" because car-
bonised soft internal organs such as the digestive tract, nerves,
and muscles can be seen through the transparent cuticle. Both
dorsal and ventral surfaces, in addition to the soft internal
organs, are sometimes preserved in a single specimen,
presenting complicated morphological puzzles. Thus, the
Tshekarda material offers a rare view of the morphology of
Paleozoic insects and an opportunity to acquire data for deter-
mining the ground plans of a diversity of organs.
The type locality and geological horizon of the Diaphanop-
terodea described here are as follows: left bank of the Silva
River, near the village of Tshekarda, Suksunsky district, in the
region of Perm, Middle Ural Mountains, Russia; Lower Per-
mian (Kungurian), Koshelevskaya series, Uppermost Irensky
horizon.
The Diaphanopterodea and the pterygote ground plan
Of all Paleozoic insects, the specimens of U. maculuta from
the Urals contribute the largest number of clues to ancestral
pterygote characters that can be used to reconstruct morpho-
 230 CAN. J. ZOOL.
logical ground plans (Kukalovsi-Peck 1983, 1985, 1991,
1992).
The Biaphanopterodea are the only pterygote order that
retains the ground-plan pterygote wing articulation (KukalovB-
Peck 1983, 1991; Kukalovsi-Peck and Brauckmann 1990).
Theoretically this does not necessarily imply that most or any
of their other morphological structures are equally primitive,
yet the potential for the Diaphanopterodea to have retained
other basic character states is clearly high. In 1976, Prof.
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B. B. Rohdendorf, then the head of the Laboratory of Arthro-
pods, Paleontological Institute of the Academy of Sciences,
USSR, sent the entire Tshekarh collection of Diaphanop-
terodea to J. Kukdovsi-Peck, who studied all the then-
available fossil evidence pertaining to hexapod, insectan, and
pterygote morphological ground plans. The morphological
structure of diaphanopterid heads and bodies was analysed, com-
pared, and interpreted over a period of 12 years (Kukalovi-Peck
1978, 1983, 1985, 1991; Shear and Kukdovsi-Peck 1990;
KukalovB-Peck and Brauckmann 1990). The evidence became
integrated over the years into many complex morphological
arguments. The detailed documentation of the heads and bodies
is presented in the accompanying paper (Kukalovsi-Peck 1992).
The purpose of this paper is to describe the best preserved
Uralian Diaphanopterodea, Wralin mculata n.sp. and Uralia
For personal use only.
shnrovi n. sp. (Parelmoidae) and Pamralia rohdendorj? n .sp.
and Paruralia carpenteri n. sp. (Paruraliidae) , and present
phylogenetic considerations (based mostly on wing venation,
as is customary for Paleozoic insects).
Systematics
Paleoptera
SUPERORDER Paledictyopteroidea
ORDER Diaphanopterodea Hand1.irsch, 1906 (nomen
trmshtum Rohdertdorf, 4%2)-
- - - - - - - - -
FAMILY Parelrnoidae Rohdendorf, 1962
OCCURRENCE: Lower Permian of North America and
Europe; warm-temperate climatic zone.
Wing shape and wing venation provide the principal distinc-
tive characters for the taxonomic evaluations of Paleozoic
insects. However, owing to the scarcity of fossils, their patchy
occurrence, and incomplete preservation, the strength and
value of venational characters af different taxonomic levels are
often uncertain and have to be adjusted when additional or
better preserved specimens -are described.
The diaphanopterid family Elmoidae was established by
Tillyard (1937) for Elmoa (Lower Permian of Oklahoma).
Carpenter (1943, 1947) added two genera, Parelmoa and
Pseudelmoa (Lower Permian of Kansas), and offered a rather
broad concept for the Elmoidae, with emphasis on the type of
the typical diaphanopterid brace MA-RP. This brace is
expressed in the Biaphanopterodea either as a short crossvein
connecting MA with RP, or as MA being adjacent to or fused
with RP for a shorter or longer distance, or MA may fuse with
R and subsequently with RP and diverge from RP. The Elmoi-
dae were defined as having a primitive, crossvein brace.
Later, Rohdendorf (1942) transferred Parelmoa and Pseudel-
msa from Elmoidae to a new family Parelmoidae, based on the
following differences: longer ScP and larger anal area with
richly branched anal veins. Kukalovsi-Peck (1974) followed
Carpenter and added to Elrnoidae six more genera from the
Lower Permian of Czechoslovakia (Elmodiapha, Proto-
diapha, Diapha , Permotliapha , Stenodiapha, and Para-
VOL. 70, 1992
diapha). The European genera had relatively long ScP and
richly branched anal veins (as in Parelrnoidae), but combined
with broadly varied MA -RP brace (crossvein, coalescence,
or fusion).
By far the best documented diaphanopterid genus is Uralia
as described here. It shows a very close similarity to Czechos-
lovakian genera by bearing long ScP and richly branched anal
branches (as ,in Parelmoidae) combined with braces (fusions)
either betweim MA and RP (U.macuhta, Figs. 2, 3) or
between MA, the end of R, and RP ( Urnlia sharovi, Fig. 5).
The implication is that the veinal characters of Parelmoidae
occur consistently in North American and Central and East
European genera, whereas the MA-RP brace shows an
increase in specialisation from west to east. Clearly, the
MA-RP brace is not stable enough to be taken as a family
character and can be used only as a generic or a specific
character. The following venational features are proposed as
being important for distingishing the Elmoidae and Parelmoi-
dae at the family level:
l . Elmoidae Tillyard, 1937 (Lower Permian) is a monotypic
family differing from Parelmoidae by having shorter ScP end-
ing before midwing, wings gradually narrowed basally, and
narrow anal area containing long AA1+2 and very short
AA3 +4, while AP is reduced; the MA -RP brace is formed
by a crossvein (plesiomorphy within the Diaphanopterodea),
and its variability within the Elmoidae is not known.
2. Parelmoidae Rohdendorf, 1962 is a valid family and con-
tains diaphanopterids with more or less oval wings bearing a
relatively broad anal area with AA1+2 long and often
+
branched, AA3 4 often branched, and AP1+ 2, and some-
times also short AP3 +4, present; MA-RP brace varies
broadly from a simple crossvein to a temporary coalescence
and fusion; MA may fuse temporarily not only with RP but
also with the-end of R.
The family Parelmoidae shares the following features with
the Elmoidae and Diaphanopteridae: highly specialised sectors
CUA and CUPrunning basally parallel and very close to each
other and connected by at least one short, prominent crossvein
(Figs. 2 -5); CuA is either simple or with a small terminal
fork; and RP branches are simple (all synapomorphies). The
Lower Permian Parelmoidae is possibly the sister-group of the
Lower Permian Elmoidae, whereas the Upper Carboniferous
(Stephanian; ca. 30 million years older) Biaphanopteridae,
with MP and CUPbranched and the parallelism of Cu sectors
less pronounced, is more primitive and may have included the
ancestor.
Uralia n. gen .
TYPE SPECIES: Uralia maculata n. sp.
ETYMOLOGY: From the Ural Mountains, Russia.
OCCURRENCE: Lower Permian (Kungurian) , Tshekarda,
Ural Mountains, Russia.
SPECIES:Uralia maculata n .sp .; Uralia ssharovi n .sp .
The name Uralia maculata and specimens 1700/496,
1700/497, and 1700/498 were mentioned previously by
Sharov (1973, p. 52) in an article on the mouthparts and feed-
ing strategies of the Paleodictyoptera. We recognise Nos.
1700/496 and 1700/497 as U. maculata, but No. 1700/498
belongs to the Protelytroptera. We consider it expedient to use
the name chosen by Sharov, but since no description or
differentiation has ever been given, the taxa are described here
for the first time.
 A N D SINICHENKOVA
DIAGNOSIS: Uralia n.gen. differs from the closely related
genus Diapha Kukalova-Peck, 1974 from Czechoslovakia by
having more numerous ScP twigs (over 30 in the forewings),
the Cu stem not flattened and ribbonlike, AA1+2 less
branched (terminal fork only), AP more branched (4-6
branches), the MA-RP brace slightly longer and more
pronounced, and wings dark colored, with scattered oval to
rounded light spots.
Description
Body
Mid-sized (wing length about 15 mm) primitive Diaphanop-
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terodea. Head relatively small, with large protruding eyes;
antennae long, filiform, with four robust basal segments
(plesiomorphy within the Pterygota); rostrum strong and rela-
tively long, mandibles with serrated inner margins (probably
primitive within the Paleodictyopteroidea); prothoracic wing-
lets fused with protergum irito large prothoracic shield (apo-
morphy) with prothoracic winglets distinctly delimited by
sutures, crossed by minute transverse ridges (apomorphy);
thorax robust; meso- and meta-thorax almost of the same size
(plesiomorphy) . Legs short and robust, with patella separated
by suture from tibia; small oblique basitarsus and 2 long eutar-
sal subsegments; post-tarsus pointed, post-tarsal claws strong
(all these are plesiomorphies within the Hexapoda). Forelegs
articulated close to head (synapomorphy of Paleodictyop-
teroidea). Abdomen with 10 full segments. Abdominal seg-
For personal use only.
ments 1-8 each with a pair of fully segmented leglets
frequently ending in tiny double claws (plesiomorphy within
the Hexapoda); abdominal pleuron composed of a separate
subcoxal, coxal, and trochanteral plate (plesiomorphy within
the Insecta); cerci very long and robust, present in both sexes
(apomorphy of the Paleoptera).
Males-Abdomen broadly attached basally, tapering abruptly
from segment 6. Folded wings longer than abdomen with geni-
talia. Genitalia on segment 9 consist of coxopodite projecting
dorsally to form a narrow, medially fused bridge; 8-segmented
leglike gonostyli ending in weak double claws; obliquely annu-
lated, strongly sclerotised, and deeply ducted penes; weakly
sclerotised, transversely annulated gonapophyses have inner
margins irregularly incised with mirror symmetry. This type
of male genitalia is close to the pterygote ground plan.
Females-Abdomen broadly attached basally, tapering very
gradually. Folded wings slightly shorter than body. Tergum 9
as long as terga 7 and 8 combined, with two large bulges
where the muscles operating the ovipositor are inserted (apo-
morphy of Paleodictyopteroidea). Pleuron of segment 8 rela-
tively narrow, with subcostal plate reinforced by oblique ridge
and coxo-trochanteral plate fused. Genitalia on segment 9 have
gonangulum attached to tergum 9 (apomorphy of the Paleo-
dictyopteroidea); broad gonocoxite covering about two-thirds
of valve 9; gonostyli leglike, with 8 segments, ending in
double claws; gonostyli long and relatively thin, densely
covered by long bristles (apomorphy); ovipositor valves
strong, gently curved, with simple, oblique, cutting ridges in
the distal half; valves protruding very little beyond the end of
abdomen. Segment 10 much shorter than pregenital segments.
This type of ovipositor is close to the pterygote ground plan.
Wings
Forewings almost oval, hind wings slightly shorter, with
straighter anterior margin and evenly arched posterior margin;
wings dark with scattered light oval spots. Costal area very
long with about 30 serial twigs from ScP and RA, several
23 1
twigs near wing base pointing towards body (apomorphy of
Diaphanopterodea). ScP ending on RA shortly beyond mid-
wing; R and M stems fused basally (apomorphy of the
Diaphanopterodea); Cu stem running close to parallel with
R & M; M diverges from R very close to where Cu divides
into CuA and CUP; RP diverges from RA shortly before mid-
wing; RP1+2 with 3 branches, RP3+4 simple; M divided
close to division of R; MA simple, temporarily fused near base
with RP, or with R and RP; MP with simple fork shorter than
MP stem; CuA adjacent near base rather shortly to M; CuA
and CUPsimple, close to each other basally and connected by
an oblique crossvein; AA 1 + 2 and AA3 + 4 with short terminal
forks; AP 4- to 6-branched. Anal brace formed by inconspicu-
ous sclerotisation of membrane basally in anal and cubital area
(plesiomorphy within the Pterygota).
Wing folding-Roofli ke , wings not overlapping at midline
(plesiomorphy within the Pterygota).
Uralia maculata n .sp.
Figs. 1-3
HOLOTYPE: Specimen 170013308, deposited in the Paleonto-
logical Institute, Academy of Sciences, Russia, Moscow.
Almost complete insect with folded wings and well-preserved
male genitalia.
ETYMOLOGY: From macula, Latin for "spot."
OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Ural Mountains. Russia.
Description
Rostrum slightly longer than head; inner mandibular mar-
gins serrated. Pronotum slightly longer than the head without
rostrum. Thoracic femora slightly longer, and tarsi slightly
shorter, than tibiae. Abdominal leglets usually pointed and
shorter than 112 length of segment (but longer close to geni-
talia). Cerci almost 2.5 x longer than body, with short trans-
versely striped annuli bearing short dense hairs and somewhat
longer lateral hairs. About 7- 10 oval spots in wing mem-
brane; apex dark. About 30 twigs in costal area; some of ScP
branches near base in the forewing forked; RP 4- to
5-branched; AP1+2 in forewing 3-branched.
Dimensions
Wing length 13.5- 18.0 mm; maximum width 4.6 -5.2 mm.
Wingspread about 40 mm. Body: head length (without ros-
trum) 2.0-2.8 mm; thorax length 6.0-7.2 mm; abdomen
length (without appendages) 12.0 - 19.0 mm. Legs: forelegs,
femur ca. 2 mm; patello-tibia ca. 1.7 mm; tarsus ca. 1.5 mm;
hind leg, femur ca. 2 mm; patello-tibia ca. 2.5 mm; tarsus ca.
2 mm. Ovipositor 6.8-7.0 mm long. Cerci ca. 57 mm long.
MATERIAL
Holotype and paratypes Nos. 17001415, 17001483- 17001490,
17001492 - 17001497, 170013300, 170013303- 170013305,
and 170013307.
Relationships
Uralia maculata n.sp. differs from Uralia sharovi n.sp. in
having fewer light spots in the wing membrane, fewer twigs
in the costal area, shorter brace between MA and RP (fusion)
and between CuA and M (coalescence), and fewer AP
branches.
 232 CAN. J . ZOOL. VOL. 70, 1992
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FIGS. 1-6. Diaphanopterodea, Parelmoidae, Lower Permian (Kungurian) of Tshekarda, Ural Mountains, Russia. Fig. 1. Uralia maculata
n. sp., holotype, No. 170013308, male, wing membrane dark with light spots; body length ca. 17 mm; forewing length 14.3 mm. Fig. 2. Uralia
maculata n.sp., forewing, based on the holotype and paratype 17001494. Length 14.3 mm. Fig. 3. Uralia maculata n.sp., hind wing, based
on the holotype and paratype 170013307,484. Length 12.5 mm. Figs. 4 -6. Uralia sharmi n.sp., holotype, No. 17001491. Fig. 4. Rooflike
position of wings at rest. Fig. 5. Forewing, dark with circular spots. Length of forewing fragment 13 mm. Fig. 6. Photograph of the holotype.
Length of left forewing fragment 13 mm, total length of wing ca. 15 mm. All originals.

Uralia sharovi n. sp. right forewing crossed with hind wing.

Figs. 4-6 ETYMOLOGY: The species is named in honor of Dr. A. G.
Sharov, an outstanding entomologist and the collector of the
HOLOTYPE: Specimen 17001491, deposited in the Paleonto- Diaphanopterodea at ~shekarda.-
logical Institute, Academy of Sciences, Russia, Moscow. OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Mesothorax, left forewing with clearly preserved venation, Ural Mountains, Russia.
 KUKALOVA-PECK A N D SINICHENKOVA
Description
Fore wing
About 25 scattered oval to round spots in wing membrane,
including apical area. Costal area with about 20 simple serial
twigs; MA fused shortly with RA before fusing with RP; CuA
adjacent near base temporarily to M; AP1+2, AA1+2,
AA3+4, and AP1+2 terminally forked, AP3+4 long,
4-branched.
DIMENSIONS
Length of wing fragment 13.0 mm, presumed total length
about 15 mm; maximum width 4.5 mm.
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Material
Holotype only.
Relationships
Uralia sharovi n.sp. differs from U. maculata n.sp. in hav-
ing more (ca. 25) light spots in the wing membrane, fewer
(ca. 20) twigs in the costal area, MA fused with the end of R
and with RP, CuA coalesced with M for a greater distance,
and AP with more (about 6) branches.
FAMILYParuraliidae n. fam.
TYPE GENUS: Paruralia n.gen.
OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Ural Mountains, Russia.
For personal use only.
GENERA: Paruralia n.gen.
Description
Body
Mid-sized diaphanopterids. Head and eyes relatively large,
almost as broad as prothorax; rostrum strong and relatively
long; prothoracic shield not known; femora short and robust,
patello-tibiae short and slender; basitarsus oblique, small,
tarsus with 4 subsegments, relatively long and thin. Abdomen
10-segmented. Folded wings in females slightly shorter than
body. Tergum 9 about as long as terga 7 and 8 combined, with
two large muscle-related bulges. Female genitalia have gonan-
gulum attached to tergum 9 and gonocoxite flap small, cover-
ing only the upper part of valve 9; gonostyli articulated distally
on gonocoxite, not preserved; valves 8 gently curved, valves
9 almost straight; valves 8 and 9 with fine, cross-hatched
ridges with posteriorly pointing denticles creating a coarse
file. Pleuron of segment 8 as in U. maculata n.sp. Segment 10
almost as long as segment 7; ovipositor valves protruding con-
siderably beyond segment 10.
Wings
Forewings with narrow, tapering basal third, dark with
rounded spots mostly combined into ca. 7 oblique stripes.
Pterostigma sometimes present. Costal area narrow and
moderately long, with ca. 15 serial twigs from ScP and only
up to 6 from RA; several twigs near base pointing towards
body; ScP ending on RA shortly beyond midwing; R and M
stems fused basally, Cu stem running close and in parallel to
R & M; M diverged from R beyond division of Cu; RP
diverged from RA well before midwing; RP1+ 2 with anteri-
orly turned tip, ca. 5-branched, RP3+4 3- to 5-branched; M
divided well before R divides; MA simple, fused shortly with
RP; MP 5- to 6-branched, MP branches longer than MP stem;
CuA adjacent basally very shortly to M; CuA and CUP with
short terminal forks or simple, parallel, but not close to each
other basally, not connected with a conspicuous short cross-
vein; AA1+2 forked, AA3+4 and AP1+2 with terminal
233
fork, AP3 + 4 very short, simple. Anal brace formed by an
oblique sclerotised bar basally between AA 1 + 2 and Cu stem
and a sclerotised membrane.
Wing folding
More or less flat over abdomen, wings partly overlapping at
midline (apomorphy, possibly of the Paruraliidae) .
Relationships
The new family Paruraliidae does not have specialised
veinal sectors CuA and C U Prunning basally very close to each
other as in the Diaphanopteridae, Elmoidae, and Parelmoidae.
Two probable synapomorphies are shared with the Martyn-
oviidae Tillyard, 1932: a broad area between AA1+2 and the
Cu stem near the wing base (the site of ridgelike sclerotisation
serving as an anal brace in Paruralia; Figs. 8 - lo), and a dis-
tinctively shaped pterostigma extending posteriorly beyond
RA (preserved in Paruralia rohdendorj7; an identically shaped
pterostigma preserved in Martvnovia and Eumartynovia;
Carpenter 1963). The Paruraliidae are more primitive than the
Martynoviidae in having RP3+4 3- to 5-branched, MP 5- to
6-branched, and most anal veins branched, and may be ances-
tors or near ancestors of the Martynoviidae.
Paruralia n.gen.
TYPE SPECIES:Paruralia rohdendor- n.sp.
ETYMOLOGY: From p a r a (Latin for "next to" and Uralia.
OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Ural Mountains, Russia.
SPECIES:Paruralia rohdendor- n.sp., Paruralia carpen-
teri n.sp.
Description
By monotypy, identical with that of the family
Paruralia rohdendorfi n.sp.
Figs. 7-9
HOLOTYPE: Species 170013304, deposited in the Paleonto-
logical Institute, Academy of Sciences, Russia, Moscow.
Complete insect with folded wings; head, fore, and middle
legs and an ovipositor in lateral view partly preserved.
ETYMOLOGY: Named in honor of Prof. B. B. Rohdendorf,
the founder of the only paleoentomological laboratory in exis-
tence, and a distinguished entomologist.
OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Ural Mountains, Russia. .
DIAGNOSIS: Paruralia rohdendor- n.sp. differs from P. car-
penteri n.sp. in having wings narrower in the basal quarter,
pterostigma present, light spots less numerous and more
merged together in stripes, more twigs in the costal area
(ca. 24), shorter fusion (brace) between MA and RP, and a
relatively large number of crossveins.
Description
Rostrum slightly longer than head; pronotum almost as long
as head without rostrum. Prothoracic femora slightly shorter
than patello-tibiae; basitarsus short, oblique; tarsus longer
than patello-tibia, thin; 4 tarsal subsegments about equally
long. Forewings very narrow in basal quarter. Light spots in
wing membrane merging into about 7 oblique stripes. Costal
area with about 24 twigs from ScP and RA.
Head length without rostrum 2.2 mm; thorax length
5.6 mm; forewing length 20 mm, maximum width 5.2 mm;
wingspread about 42 mm. Right foreleg: femur ca. 2.0 mm;
patello-tibia ca. 1.9 mm; tarsus ca. 2.4 mm.
 CAN. I . ZOOL. VOL. 70. 1992
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FIGS.7 - 10. Diaphanopterodea, Paruraliidae n.fam., Lower Permian (Kungurian) of Tshekarda, Ural Mountains, Russia. Figs. 7 -9. Paruralia
rohdendorji n.sp. Fig. 7. Holotype, No. 170013309; wing membrane dark with light stripes. Total length ca. 28 mm. Fig. 8. Holotype, showing
rooflike position of wings at rest. Fig. 9. Forewing with oversized pterostigma. Length 20 mm. Fig. 10. Pctruralia carpenteri n.sp., holotype,
No. 170011873, light spots partly merged into stripes. Forewing length 13 mm. All originals.

Material Relationships between diaphanopterid families

Only the holotype is known.
Paruralia carpenteri n .sp.
Fig. 10
HOLOTYPE: Specimen 170011873 deposited in the Paleonto-
logical Institute, Academy of Sciences, Russia, Moscow. A
single forewing, well preserved.
ETYMOLOGY: In honor of Prof. F. M. Carpenter, a distin-
guished entomologist whose studies have advanced the under-
standing of Diaphanopterodea and other Paleozoic insects.
OCCURRENCE: Lower Permian (Kungurian), Tshekarda,
Ural Mountains, Russia.
Diagnosis and description
Paruralia carpenteri n.sp. differs from the related P. roh-
dendor- n.sp. in that its forewings are less narrowed basally,
with more numerous and mostly rounded light spots partly
merged into stripes, pterostigma absent, apex pointed, only
about 18 twigs in costal area, longer fusion (brace) between
MA and RP, and few crossveins.
DIMENSIONS: Forewing length 13 mm, maximum width
4 mm.
MATERIAL: Only the holotype is known.
The order Diaphanopterodea was based by Rohdendorf
(1962) on the combination of paleopterous wing venation and
the ability to flex the wings backwards over the abdomen while
resting, a character unique to this order within the Paleoptera.
The body structures were little known at first, but during the
last 12 years a wealth of information has been added on wing
venation, wing articulation and folding, the detailed structure
of the mouthparts, male and female genitalia, prothorax, and
abdominal pleuron, and leg segmentation, mainly because of
the remarkable degree of preservation of the specimens of
Uralia (Kukalova-Peck 1983, 1985, 1987, 1991, 1992). Some
of these features can be used to reconstruct the phylogeny, as
shown below. The presently known families seem to fall into
the following three groups:
1. Diaphanopteridae (ancestors), Elmoidae, and Parelmoidae
(Carpenter 1947, 1963; this study) (descendents and (or) pos-
sible sister-groups) share Cu sectors that are basally parallel
and close to each other, nearly simple CUA, and simple RP
branches (synapomorphies). Ovipositor of the cutting type,
with simple transverse ridges (plesiomorphy within the Ptery-
gota). Prothoracic shield well developed but with wings still
recognisable, delimited by sutures (possible sy napomorphy) ;
3 tarsal segments (plesiomorphy of the Pterygota) (body struc-
tures known only in the Parelmoidae: Uralia).
 KUKALOVA-PECK A N D SLNICHENKOVA
2. Prochoropteridae (primitive; Fig. 6 in Kukalova-Peck and
Brauckmann 1990), Paruraliidae (moderately derived), Mar-
tynoviidae, and the asthenohymenids (including Astheno-
hymenidae, Permohymenidae, and several undescribed Lower
Permian families from Obora, Czechoslovakia; J. Ku kalova-
Peck, unpublished data) (highly derived; possibly sister-group
lineages); share relatively narrow to very narrow costal area,
broad area between A A and Cu near wing base, and increas-
ingly longer braces, coalescence between CuA and M , and
fusion between M A and R A - R P (all synapomorphies).
Ovipositor known in Paruraliidae and Prochoropteridae (in
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Euchoroptera longipvnnis Carpenter, 1933) (Kukalova-Peck
1992) and in the asthenohymenids from the Lower Permian of
Czechoslovakia (J. Kukalova-Peck, unpublished data) is
cross-hatched with denticles and operates like a file (apomor-
phy). Prothoracic shield either not formed and prothoracic
wings present. pointed, protruding laterally (plesiomorphy
within the diaphanopterodea: in Prochoropteridae), o r fully
formed (in asthenohymenids ; J . Kukalova-Pec k, unpublished
data); 5 tarsal segments (? synapomorphy; known in Paruralii-
dae and in the asthenohymenids).
3. Namurodiaphidae Kukalova-Peck and Brauckmann, 1990,
the oldest family of Diaphanopterodea from the early Upper
Carboniferous (Bashkirian; Namurian B), occurs with the
oldest fossilised Pterygota. Namurodiaphidae has the most
primitive venation from which the venation of group 1 as well
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as group 2 can be derived; ovipositor identical with that of
group I (symplesiomorphy). Prothoracic shield not formed,
prothoracic wings present, pointed, protruding laterally as in
group 2 (symplesiomorphy); 3 tarsal segments, as in group 1
(symplesiomorphy) (body structures known only from
Numurodiuphu sippelorurn, holotype).
In the past, an attempt was made by Rohdendorf (1962) to
subdivide the Diaphanopterodea into two suborders: the
Metadiaphanopterodea with the Asthenohymenidae, and the
Eudiaphanopterodea containing the remaining families (fol-
lowing Rohdendorf 1962, and emendations of diaphanopterid
families by Carpenter 1963). The evidence summarised above
shows that this subdivision is not natural and that the Diaphan-
opterodea probably should not be subdivided into suborders,
given the present level of knowledge.
Acknowledgements
This account would not have been possible without the gener-
ous loan of the unique fossil material by Dr. B. B. Rohdendorf
(Laboratory of Fossil Arthropods, Paleontological Institute,
Academy of Sciences, Moscow) to the first author. Dr. A . P .
Rasnitsyn, the present head of the laboratory, showed a deep
understanding of and patience with the great amount of time
required to make the detailed morphological cross-comparisons.
T o both of these paleoentomologists we offer our sincere grati-
tude. Help of various kinds was given and photography was
expertly performed by Mr. L. E. C . Ling (Carleton Univer-
sity, Ottawa). The project was supported by operating grants
from the Natural Sciences and Engineering Research Council
of Canada. Facilities and work space were provided by the
Department of Earth Sciences, Carleton University, Ottawa.
235
Carpenter, F. M. 1943. The Lower Permian insects of Kansas. Part
9. The orders Neuroptera, Raphidiodea, Caloneurodea and Protor-
thoptera (Probnisidae) with additional Protodonata and Mega-
secoptera. Proc. Am. Acad. Arts Sci. 75(2): 55 -84.
Carpenter, F. M. 1947. Lower Permian insects from Oklahoma. Part
1. Introduction and the orders Megasecoptera, Protodonata, and
Odonata. Proc. Am. Acad. Arts Sci. 76(2): 25 -54.
Carpenter, F. M. 1963. Studies on Carboniferous insects from Com-
mentry, France. Part V. The genus Diaphanopteru and the order
Diaphanopterodea. Psyche (Camb.) , 70: 240 - 256.
Kalugina, N. S. 1980. The insects in the ecosystems of the past.
In The historical development of the class Insecta. Edited by B. B.
Rohdendorf and A. P. Rasnitsyn. (In Russian.) Tr. Paleontol. Inst.
Akad. Nauk SSSR. 175: 224-239.
Kukalova-Peck, J . 1974. Wing folding in the Paleozoic insect order
Diaphanopterodea (Paleoptera), with a description of new
representatives of the family Elmoidae. Psyche (Camb.), 81:
315-333.
Kukalovh-Peck, J . 1978. Origin and evolution of insect wings and
their relation to metamorphosis, as documented by the fossil
record. J . Morphol. 156: 53- 126.
Kukalovi-Peck, J . 1983. Origin of the insect wing and wing articula-
tion from the arthropodan leg. Can. J. Zool. 61: 1618 - 1669.
Kukalova-Peck. J . 1985. Ephemeroid wing venation based upon new
gigantic Carboniferous maytlies and basic morphology, phylogeny,
and metamorphosis of pterygote insects (Insecta, Ephemerida).
Can. J . Zool. 63: 933-955.
Kukalova-Peck, J . 1991. Fossil history and the evolution of hexapod
structures. In The insects of Australia. 2nd ed. Edited by I. D.
Naumann and CSIRO. Melbourne University Press, Melbourne.
pp. 141-179.
Kukalova-Peck, J . 1992. The "Uniramia" do not exist: the ground
plan of the Pterygota as revealed by Permian Diaphanopterodea
from Russia (Insecta: Paleodictyopteroidea). Can. J . Zool. 70:
236 -255.
Kukalova-Peck, J . , and Brauckmann, C. 1990. Wing folding in
pterygote insects, and the oldest Diaphanopterodea from the early
Late Carboniferous of West Germany. Can. J . Zool. 68: 1104-
1111.
Martynov, A. V. 1938. Review of localities of fossil insects in USSR.
Tr. Paleontol. Inst. Akad. Nauk SSSR, 7(3): 7-28.
Rasnitsyn. A. P. 1980. Superorder Diaphanopteridea Handlirsch,
1906. In The historical development of the class Insecta. Edited by
B. B. Rohdendorf and A. P. Rasnitsyn. (In Russian.) Tr.
Paleontol. Inst. Akad. Nauk SSSR, 175: 49-52.
Rohdendorf, B. B. (Editor). 1962. Fundamentals of paleontology.
Tracheate and chelicerate arthropods. (In Russian.) Acad. Nauk.
SSSR, Moscow. pp. 1 -560.
Sharov, A. G. 1973. The morophological characters and the way of
life of Palaeodictyoptera. (In Russian.) In Lectures in memory of
N. A. Cholodkovskii, 1971. Academiya Nauk. Moscow. pp.
49-63.
Shear, W. A., and Kukalova-Peck, J . 1990. The ecology of Paleozoic
terrestrial arthropods: the fossil evidence. Can. J . Zool. 68:
1807-1834.
Sinichenkova. N. D. 1987. The historical development of stonetlies.
(In Russian.) Tr. Paleontol. Inst. Akad. Nauk SSSR, 221: 1 - 143.
Tillyard, R. J . 1937. Kansas Permian insects. Part 17. The order
Megasecoptera and additions to the Palaeodictyoptera, Odonata,
Protoperlaria, Copeognatha, and Neuroptera. Am. J . Sci. 33:
81 - 110.