Nearly all of the fossils taken
from the Kansas formations were of winged insects
(Subclass Pterygota). However, one primitively unwinged insect fossil
(Subclass Apterygota) was described in 1978. Two subdivisions are often used by paleontologists to further differentiate
the winged insects:
The Palaeoptera (literally ancient-winged)
and Neoptera (modern-winged). The Neoptera can fold their wings down flat
against the dorsal surface of their abdomen (like
the bees, beetles, and flies of today).
Most of the paleopterous insects had to hold
their wings out at right angles to the body (like
the dragonflies or the mayflies), although one order
(the Diaphanopterodea) did have a simpler mechanism
that allowed them to fold their wings back in a similar
manner to the Neoptera.
Checklist of Permian Insects of Elmo, Kansas:
Compiled by Roy J. Beckemeyer.
This list contains only the most recent names.
Many of the species names used in the literature
were later synonymized. Contact the author for a version of this list containing all synonyms.
Higher order taxonomy is per Ref. 19.
Key:
No indentation: Order (Higher taxon) Eras.
Notes;
-One tab: Suborder;
--Two tab: Family Author Date described;
---- Four tabs: Genus Author Date described;
------ Six tabs: species Author Date described.
|
|
| Fig.
7. L. sharovi
after Durden 1978 (Ref. 25) |
Archaeognatha (Apterygota) U.Carb.-Hol.The bristletails are primitive
wingless insects with a long "caudal process"
or tail, but without cerci. The thoracic segments
look very much like the abdominal segments dorsally. One
family, 1 genus, 1 species.
- Dasyleptidae Sharov 1957
-- Lepidodasypus
Durden 1978
---- sharovi
Durden 1978 (Fig. 7)
|
|
| Fig.
8. P. permianum
after Tillyard, 1932 (Ref. 62) |
Ephemeroptera (Pterygota: Palaeoptera) U.Carb.-Hol.Mayflies are aquatic
insects. This is the only palaeopterous order that
has existed from the Upper Carboniferous to the present.
Current adult mayflies do not have functional mouthparts
and do not eat; fossil mayfly adults had chewing mouthparts.
The hind wings of modern forms are reduced or absent;
in fossil forms the fore and hind wings are almost
the same size. Two families, 2 genera, 8 species.
- Misthodotidae Tillyard 1932
-- Misthodotes
Sellards 1909
---- obtusus
(Sellards) 1909 (Dromeus)
---- biguttatus
Tillyard 1932
---- ovatus
Tillyard 1932
- Protereismatidae Lameere 1917
-- Protereisma
Sellards 1907
---- permianum
Sellards 1907 (Fig's. 8, 20)
---- elongatum
(Sellards) 1907 (Bantiska)
---- arcuatum
(Sellards) 1907 (Rekter)
---- latum
Sellards 1907
---- insigne
Tillyard 1932
Palaeodisctyoptera (Pterygota:
Palaeoptera) U.Carb.-Perm. These insects had winglets
on their prothorax.
They held their wings open much like modern
dragonflies. They
had sucking mouthparts as did the next two orders,
Megasecoptera and Diaphonopterodea, which together
with the Palaeodictyoptera make up the palaeodictyopterid
group. It
is thought that these insects fed on plant juices
or on the semi-liquid endosperm of Paleozoic plants.
Three families, 4 genera, 4 species.
- Calvertiellidae Martynov 1931
-- Calvertiella
Tillyard 1925
---- permiana
Tillyard 1925
- Elmoboriidae Carpenter 1976
-- Elmoboria
Carpenter 1976
---- piperi Carpenter 1976
- Spilapteridae Handlirsch 1906
-- Dunbaria
Tillyard in Dunbar & Tillyard 1924
---- fasciipennis
Tillyard in Dunbar & Tillyard 1924 (Pictured
on front & back covers.)
- Uncertain Family
-- Kansasia
Tillyard 1937
---- pulchra
Tillyard 1937
Megasecoptera (Pterygota: Palaeoptera)
U.Carb.-Perm. A group very similar to the previous
order, but with the fore and hind wings nearly equal
in size, and noticeably long and slender.
Wing bases often very slender and reminiscent
of those of modern damselflies. One family, 2 genera,
5 species.
|
|
| Fig.
9. P. readi
after Carpenter 1933 (Ref. 9) |
-
Protohymenidae Tillyard 1924
-- Protohymen Tillyard 1924
---- permianus Tillyard 1924 (Photo on back
cover)
---- schucherti Tillyard 1926
---- elongatus Carpenter 1930
---- readi Carpenter 1933 (Fig. 9)
---- tenuis Carpenter 1938
-- Permohymen Tillyard 1924
---- schucherti Tillyard 1924
Diaphanopterodea
(Pterygota: Palaeoptera) U.Carb.-Perm. Similar
to the previous order, but have the ability to fold
their wings back along the abdomen.
Three families, 3 genera, 6 species.
|
|
| Fig.
10. A. dunbari after Carpenter 1939 (Ref. 11) |
-
Asthenohymenidae Tillyard 1924
-- Asthenohymen Tillyard 1924
---- dunbari Tillyard 1924 (Fig. 10)
---- affinis Tillyard 1926
---- pusillus Tillyard 1926
- Elmoidae Tillyard 1937
-- Elmoa Tillyard 1937
---- trisecta Tillyard 1937
- Martynoviidae Tillyard 1932
-- Martynovia Tillyard 1932
---- insignis Tillyard 1932
---- protohymeniodes (Tillyard) 1932
(Martynoviella)
|
|
| Fig.
11. A full-sized image of a hind-wing of
Megatypus schucherti Tillyard
1924. This
fossil is in the Kansas State University Entomology
Dept. collection. Scanned by Roy Beckemeyer with the permission
of Dr. Ralph Charlton of KSU.
This 19.5 cm wing, found by two KSU entomology
students, Otto Wenger and Floyd Holmes in 1939,
is one of the most complete Protodonata wings
ever found – nearly all the venation is clear
and well-preserved. |
Protodonata
(Pterygota: Palaeoptera) U.Carb.-Perm.
These dragonfly-like insects were usually
large and occasionally gigantic in size.
The largest insect known is from Elmo:
Meganeuropsis permiana is estimated to have
had a wingspan of nearly 29 inches.
It was likely at the upper limits of size
for which tracheal respiration could supply enough
oxygen to its flight muscles; it probably also had
the ability to actively control its thoracic temperature,
making it one of the first endotherms (Ref. 33).
Protodonata ruled the Permian skies, preying
on flying and perching insects. Probably aquatic as are the Odonata of today.
Two families, 4 genera, 7 species.
-
Meganeuridae Handlirsch 1906
-- Meganeuropsis Carpenter 1939
---- permiana Carpenter 1939
-- Megatypus Tillyard 1925
---- schucherti Tillyard 1925 (Fig. 11)
---- ingentissimus Tillyard 1925
-- Tupus Sellards 1906
---- permianus Sellards 1906
---- readi Carpenter 1933
---- vetustus (Carpenter) 1933 (Megatypus)
- Paralogidae Handlirsch 1906
-- Oligotypus Carpenter 1931
---- tillyardi Carpenter 1931
Odonata (Pterygota: Palaeoptera)
Perm.-Hol.Dragonflies and damselflies quite similar
to modern forms were present in the Elmo fauna.
Like the protodonates, the odonates were predaceous.
It is possible that their generally smaller
size was helpful in their survival to modern times.
It has been postulated (Ref. 26) that an increase
in atmospheric oxygen level to about 35% during the
Carboniferous and Permian supported the development of very large flying insects
and that the decrease
in oxygen level at the end of the Permian contributed
to their extinction.
Two families, 4 genera, 7 species.
|
|
| Fig.
12.K. mirabilis
Tillyard 1925, Peabody Museum specimen No. 5006a,
Collector: C.O. Dunbar, 1921.Basal portion of
hind wing.Photo courtesy of Yale University. |
Archisozygoptera Handlirsch 1906
- Kennedyidae Tillyard 1925
-- Kennedya
Tillyard 1925
---- mirabilis Tillyard 1925
(Fig. 12)
---- tillyardi
Carpenter 1939
---- reducta
Capenter 1939
-- Progoneura
Carpenter 1931
---- minuta
Carpenter 1931
- Protanisoptera Carpenter 1931
-- Ditaxineuridae Tillyard 1926
---- Ditaxineura
Tillyard 1926
------ anomalostigma
Tillyard 1926
------ celulosa
Carpenter 1933
- Uncertain Suborder
-- Uncertain Family
---- Campotaxineura
Tillyard 1937
------ ephialtes
Tillyard 1937