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Volume 46, Number 1, February 2000:
The Permian Insect Fossils of Elmo, Kansas


Image - front cover of issue

ISSUE HOME PAGE

ABOUT THIS ISSUE
- about KSN
- about the author

IN THIS ISSUE
- introduction
- the elmo site as part of the Kansas-Oklahoma permian
- a short history of the discovery and study of the elmo fossils
- the insects: part 1, part 2, part 3,
- references
- back cover

SLIDESHOW
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This page was last modified:
October 15, 2003

Originally posted:
March 20, 2003


 

The Permian Insect Fossils of Elmo, Kansas
by Roy J. Beckemeyer


The Insects


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)


Image - Figure 8 P. permianum after Tillyard, 1932 (Ref. 62)
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.


Image - Figure 9 P. readi after Carpenter 1933 (Ref. 9)
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.


Image - Figure 10 A. dunbari after Carpenter 1939 (Ref. 11)
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)


Figure 11 - Full-sized image of hind-wing of Megatypus schucherti Tillyard 1924.
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.


Figure 12 - K. mirabilis Tillyard 1925
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



Next Section: the insects (continued)

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