|1. Life sized adult woman with a life sized model of a griffenfly|
People have a continuing fascination with giants. Part of the attraction of dinosaurs is their great size, the largest over 100 feet in length and 100 tons or more in weight. Many groups of plants and animals other than dinosaurs achieved gigantism sometime in their evolutionary history (lycopod plants, scorpions, rhinoceroses, and varanoid lizards among others). However, not all giants are of absolutely large size. Being crowned a “giant” depends on how large the other members of your group are. There are several forces that drive the evolution of gigantism -- anatomy, mechanical constraints, and environment to name just a few. One of the more interesting giants in the fossil record, and one with some pop culture significance, are the giant dragonflies of the Paleozoic.
Dragonflies (Order Odonata) are among the most recognizable groups of living insects. They are large, often brightly colored, active during the day, and quite visible flying around and resting in open areas. Among the insects, dragonflies have devoted human observers, the arthropod equivalent of bird watching. With at least 6000 known living species there are plenty of dragonflies to watch.
2. Large eyes, strong jaws, and powerful wings are some of the predatory hallmarks of dragonflies
All dragonflies are predators and most are highly agile aerial predators. One exception is the damselfly family Pseudostigmatidae, which hunts orb-weaving spiders and plucks them from their webs! Dragonflies can fly at speeds up to 35 miles an hour chasing prey or hover motionless over a pond. They can even fly backwards! Their large spherical eyes, covering most of their head, provide excellent vision for hunting prey --- mostly small flying insects (especially mosquitos) – by flying, catching them between their legs,and eating them on the run. Although they may live several years as an aquatic nymph, flying adults have a short life of between one and six months.
3. Dragonfly nymphs are predatory and capable of feeding on small vertebrates
The nymphs of dragonflies are also voracious predators. In some species the nymph captures, kills,and eats tadpoles and small fish! As we’ll see later, this has some interesting implications for griffenflies.
THE FOSSIL GIANTS
Dragonflies have a long evolutionary history, extending back some 300 million years to the Carboniferous Period. Technically speaking these ancient giants are not true dragonflies but rather the closest relatives of dragonflies. That need not worry us very much as the distinction is based on detailed anatomical and evolutionary analysis and is, in part, purely definitional. They are known scientifically as protodonata, meganeuridae, meganisopterans, or griffenflies. Here I’ll use griffenflies. I’m not sure of the etymology of that name, but in German "griffen" means “attack” – resulting in an appropriate nomen.
When you step on a cockroach, there is a pleasing crunchy sound, made by the compressing and cracking of the hard exoskeleton. However, the exoskeleton of insects is not always hard and not all parts are equally hard. In dragonflies it is the wings that are the most resilient body part and the part most likely to be fossilized. So, much of the fossil record of griffenflies consists of fragmentary specimens, with a few spectacular exceptions. Griffenflies darted through the Earth’s skies for some 20+ million years million years (Late Carboniferous 317 MYA – Late Permian 247 MYA) and achieved a worldwide distribution.
4. The type specimen of the griffenfly Namurotypus sippeli preserving details of the head and body
5. Restoration of Namurotypus sippeli
6. Restoration of Namurotypus sippeli
There are a few complete or nearly complete griffenflies that provide much information about the overall anatomy of the group (1, 2). They have large, robust, toothed “jaws”, large spheroidal compound eyes, and strong legs with large spines. Their wings are large, strong, and directed sideways from the body. All these are dragonfly adaptations for aerial predation and griffenflies clearly had a similar lifestyle.
8. Another unidentified, but spectacularly complete, griffenfly from France
Griffenflies achieved a wide distribution, were fairly diverse (3), and new species continue to be described every year. While most griffenflies are large, some are within the size range of modern dragonflies. However, some are truly gigantic by insect, and especially dragonfly, standards.
Meganeura monyi,the first described griffenfly, was based only a fossil wing about 12 inches long and an estimate wing span of 27 inches (including an estimated body width between the wings). At the time of its description in 1895 it was the largest insect known and remained so until 1939 (4), when Frank Carpenter described Meganeuropsis permiana from Kansas. M.permiana was based on an incomplete, but large wing, in two parts. Scaling the wing of M. monyi up to the size of M. permiana, Carpenter estimated a wingspan of 29 inches for his new species. Several years later, Carpenter described another new species of griffenfly, Meganeuropsis americana (5). This species is based on a spectacular specimen the largest complete wing ever found. He estimated the wingspan of M. americana as close to that of M. permiana. The spectacular M. americana wing is on permanent exhibit at the Museum of Comparative Zoology at Harvard University. Today, specialists consider the two species of Meganeuopsis to be the same and the correct name is M. permiana. It is this species that holds the record as the largest known insect (6). Compare this to Megaloprepus coerulatus, the living odonate with the largest wingspan --- a paltry 7.5 inches.
9. The wing of the giant Meganuropsis permiana, measuring a remarkable 13 inches!
A 29 inch wing span is damn big for an insect --- but not big enough for some. Although often repeated in the popular press, websites, and books, griffenflies NEVER achieved wingspans of FIVE or SIX feet. There is absolutely no evidence for that and such a size is almost certainly above the mechanical capabilities of the insect exoskeleton, especially for flight. Nevertheless the myth continues to be repeated as though it were a well established fact.
|10. The griffenfly Meganeuropsis permiana compared to the largest living insects. Silhouette of Lieutenant Ellen Ripley of the US spaceship Nostromo for scale|
Is it possible that the known specimens of M. permiana are not full adult size and the species grew even larger? We can answer that question definitively NO. We can do that because of the biology of dragonflies. The predatory nymph stage of dragonflies grows for months or years in lakes and streams, shedding its exoskeleton several times. However, once the nymph crawls out of the water, sheds its exoskeleton a final time, and emerges as an adult, growth ceases. So a dragonfly wing in the fossil record is of adult size. Griffenflies had a similar biology and so known wings are of fixed adult size.
This fact provides some interesting insight into griffenfly diversity. In the Permian rocks of the Lodeve Basin griffenfly wings can be grouped into five distinct size classes. Given how they grew, these must represent five different species (7).
Estimates of body length for the largest species of griffenflies, some of which are known from more than just wings, is about 17 inches. This means that the nymphs reached a length of 17 inches and must have been major predators in freshwater systems, likely feeding on amphibians as well and fish and smaller creatures.
WHY GIANT GRIFFENFLIES?
Griffenflies may have been extinct for 250,000,000 years but one might well ask why are there no equally gigantic dragonflies in today’s world? Or to turn the question around, what allowed griffenflies to attain their great size?
The Late Paleozoic was unusual in several ways. During the Late Carboniferous and Early Permian there were extensive coal swamp forests that generated tremendous amounts of oxygen as a byproduct of photosynthesis. This created a hyperoxic atmosphere, with oxygen levels in excess of modern levels. Insects lack lungs and breath through a series of tubes (trachea) that are connected to the outside. Oxygen is absorbed through the wall of these tubes by simple diffusion. With higher oxygen levels in the atmosphere the amount of the gas absorbs increases and that increased absorption would allow insects to evolve giant body size. The anatomy of griffenflies indicates very maneuverable flight capabilities which would be very demanding metabolically and require high oxygen levels. Decreasing oxygen levels in the Permian, coupled with a much more arid global aridity, could have driven these giants to extinction. Modern atmospheric oxygen levels may be too low to allow gigantism in active aerial predatory insects (8).
Of course, when griffenflies ruled the air they really ruled the air. Flying vertebrates, birds, bats, and pterosaurs, would not evolve for at least another 100,000,000 years, so the skies were free of more active fliers that might have preyed on griffenflies.
GRIFFENFLIES IN THE MODERN WORLD
“Only one scientist dared to investigate the incredible phenomena … how a simple dragonfly had become a winged monster of a species extinct for millions of years.Your flesh will crawl,Your blood will chill in terror.”
People are fascinated by giants, especially ferocious giants, hence part of the reason for the popularity of dinosaurs in movies. Other giants of the past have sometimes captured roles on the silver screen. Griffenflies appear in the 1958 black-and-white-cult-scifi-B-film Monster On The Campus. University professor Dr. Blake is studying a new specimen of the ancient fish known as the coelacanth – a “living fossil” of sorts. The writers did some background research because some of the “facts” about the coelacanth are reasonably accurate. However, the truest dialogue in the movie comes during a discussion between paleontologist Dr. Blake and a student as they stand before the coelacanth:
Dr. Blake: Do you know anything about paleontology?
Student: “Only that really attractive men study it.”
11. Meganuera (with caveman restorations behind it) getting ready to attack students and faculty in Monster On The Campus.
At one point a dragonfly (actually a damselfly) lands on the coelacanth, feeds on it, and then mutates into a giant, flying griffenfly. After terrorizing some students, Dr. Blake captures it and proclaims “This insect is of the genus Meganura, an ancestor of the present day dragonfly, supposedly extinct for millions of years.”
The quote at the beginning of this section is from the film’s trailer that can be viewed at
I won’t give away what happens to the scientist. For that you’ll have to suffer through the movie. Don't fret, I doubt “your blood will chill in terror.”
Meganeura makes continual appearances in scientific programs and museum exhibits. It appears, much better animated this time, in the BBC documentary Walking with Monsters, episode 2 in which it flies and ultimately steals a small reptile from a large spider. Meganeura also appeared the episode entitled The Bug House in ITVs six part Prehistoric Park.
|12. Prehistoric Park's host Nigel Narven with a captured Meganeura|
A very nice bit of animation with Meganeura flying around a rocky stream can be seen at
Griffenflies are also suprisingly popular in natural museum dioramas. Some examples are :
13. Restoration of flying and resting griffenflies in a coal swamp diorama in the Naturkundmuseum, Stuttgart.
|14. Meganeura in the Audubon Institute's Insectarium in New Orleans|
|15. A griffenfly model at the Denver Museum of Nature and Science|
Tania Summerell is an Australian multi-media visual artist who created a timber, acrylic perspex, plastic, and aluminum artistic rendition of Meganeura as part of her Living Fossils exhibit, which addresses issues of biotic conservation.
|16. The multi-media Meganeuropsis by Tania Sumerrell|
ROBIN TILLYARD AND THE GRIFFENFLIES
Robin Tillyard (1881-1937) was an accomplished and eminent English – Australian entomologist who worked on both living and fossil insects. He wrote extensively on the Permian insect beds of Kansas, including the griffenflies, before Chapman discovered the giants there. In addition to his many scientific papers, in 1917 he wrote The Biology of Dragonflies and in 1926 he published the authoritative The Insects of Australia and New Zealand, which remained a standard work for a half century (9).
17. Entomologist and paleo-occulist Robin Tillyard.
However, Tillyard had some mighty peculiar interests outside of insects. In modern parlance one might say he was “complicated.” He was deeply interested in the occult. After a séance with the psychic Mina Stinson (a séance which later proved to be a hoax)Tillyard became convinced of the survival of the human personality after death. In 1928 he somehow managed to get his findings published in the august scientific journal Nature. Unfortunately, his occult interests did not help him much. In 1928 the psychic Jeanne Laplace “predicted” Tillyard’s death in a motor accident and this prediction was sent to Tillyard. Tillyard’s reaction? He merely laughed at the whole affair, remarking that this type of prognostication is so seldom verified.” Maybe so, but Tillyard did indeed died in a car accident in Australia on Jan 13, 1937. Ironically, Tillyards’ daughter, Faith Evans, died on March 22, 2005 after being hit by a car (10).
This might all seem to be an illogical side to an otherwise logical scientist, except Tillyard’s occultism did seep into his paleoentomological research on at least one occasion. In their book on insect evolution, Grimaldi and Engel (2) write of Tillyard:
“It is little known that Tillyard was also a devotee of the occult, who believed that mysticism could be used to understand the lives of long vanished organisms. It has even been recounted that he once visited the Museum of Comparative Zoology [at Harvard University] with the idea of having a local Bostonian mystic bring back to life the huge Permian griffenfly specimens preserved there so that he could observe their behavior.”
Ah if paleontology were only so simple ………
GRIFFENFLIES FROM OTHER DIMENSIONS
Reincarnating the fossil griffenflies is certainly an odd idea, but not the oddest of ideas about these magnificent insects. Stace Tussel claims he had a “giant dragonfly encounter in eastern Kansas” in the 1960s (11). There on I-35 he saw a living giant dragonfly with a wingspan that could have been 5 feet. He thought this might have been the result of “experiencing a meta-dimension” or moving backwards and forwards in time. He couldn’t decide if the griffenfly exists in a parallel dimension or lives a hidden life on our planet.
And gauging from the responses, Tussel isn’t the only one to have seen those giants alive. Some thought the existence of living griffenflies in today’s world challenged evolution. Or that the beast passed through “one of the many temporal-spatial doors on our planet – some artificial, some natural.” Or that the insect “developed an invisible shield, knows how to disappear to the human look, is an interdimensional entity or high tech with insect shape that vibrates to the frequency of the camera flash.” There are more incidents and “explanations” than I can recount here, so visit Tussel’s post for some amusement --- such as that of the person who found that dragonflies speak to him, but in their own language! (11)
I live in the rural hamlet of Jensen, Utah and the bottom lands of the nearby Green River flood every Spring, some years worse than others. Regardless of the level of flooding, however, this area is famous as the mosquito factory for the state, with literally billions of those pesky West Nile Virus carrying dipterans swarming the skies. So if any readers see a living griffenfly, please let me know. We could sure use them here.
2. Griffin Chure
5 and 6. Bechly et al 2001. (ref 1)
(1) Bechly, G., Brauckmann, C., Zessin, W., and Ning, E.G.2001. New results concerning the morphology of the most ancient dragonflies (Insecta: Odonatoptera) from the Namurian of Hagen-Vorhalle (Germany).J. Zool. Syst. Evol. Research 39 (2001) 209-226.
(2) Grimaldi, D. and Engel, M.S. 2005. Evolution of the Insects. Cambridge University Press: 755 pp.
(3) Zessin, W. (2008): Überblick über die paläozoischen Libellen (Insecta, Odonatoptera). Mitteilungsblatt des Entomologischen Vereins Mecklenburg, 11. Jahrgang, Heft 1: 5-32
(4) Carpenter, F.M. 1939. The Lower Permian Insects of Kansas. Part 8: Additional Megasecoptera, Protodonata, Odonata,Homoptera, Psocoptera,Protelytroptera, Plectoptera, and Protoperlaria. Proceedings of the American Academy of Arts and Sciences 73(3): 29-70
(5) Carpenter, F.M. 1947. Lower Permian Insects from Oklahoma. Part 1.Introduction and the Orders Megasecoptera,Protodonata, and Odonata. Proceedings of the American Academy of Arts and Sciences 76(2: 25-54.
(6) Anonymous 2007. Dragonfly. The largest complete insect wing ever found. Harvard Magazine, November-December 2007. available on-line at: http://harvardmagazine.com/2007/11/dragonfly
(7)Nell, A.N., Fleckl, G., Garroustel, R., and Gand, G. 2008. The Odonatoptera of the Late Permian Lodève Basin (Insecta). Journal of Iberian Geology 34 (1) 2008: 115-122
(8)Dudley, R. 1998. Atmospheric oxygen, giant Paleozoic insects and the evolution of aerial locomotor performance. J Exp Biol. 201(Pt 8): 1043-50.
(9) John Robin Tillyard: http://en.wikipedia.org/wiki/Robert_John_Tillyard
(10) Price, L. 2005. Dr. Robin John Tillyard 1881-1937- a forgotten Australian psychical researcher. Psypioneer, An Electronic Newsletter, Vol. 1,No 19.
(11) Giant Dragonfly Encounter in Kansas: http://inter-intelligence-communications.com/?p=1231