I N S E C T F L I G H T
THE PARANOTAL THEORY
The Paranotal Theory, the generally accepted hypothesis to explain the origin of insect flight dates from 1873, when Fritz Mueller, in refuting an earlier theory, which derived wings from gills of aquatic insects, proposed that the evolutionary source of th
e wings were paranotal extensions, that is, lobes extending from the sides of the nota, which are the hard plates covering the thorax of an insect.
See fig. 1. put this in ILL: nota/singular notum)
Not only was the paranotal origin of wings simple and obvious, subsequently, fossil insects from the Paleozoic era were found, which, although already in full possession of wings, also show extensions, or lobes, of the pronotum, the first of the three thor
acic terga (dorsal plates). See fig. 2.
All that was missing now was describing the development of paranotal lobes into wings and of the muscle-driven flight mechanism. Since the function of flight is aerodynamic, ideas naturally focussed on adaptations using these lobes in traveling through air.
Such activities, with immovable lobes, or vanes, naturally suggest an origin in simpler actions involving traveling in the air, jumping and gliding. Therefore, the hinging articulation of the wings and the motive for the evolution of the flight mechanism i
s ascribed to a gradual development of refining control of movement in jumping and gliding. The flight system is here now, and it seems obvious that it must have developed in some definite manner, from a primitive, germinal source of action connected with a
erial locomotion, evolving directly, in a straight line to its present advanced state.
Explaining the origin flight mechanism relates to two parts. One is the anatomical origin of the wing, and the other, the development of the kinetic mechanism, which involves both the development of a hinged, moveable wing from the immovable, hard lobe to
its present shape, dimensions, articulations, and the muscular structure and engine that moves the wings. As for the first part, the paranotal theory offers us a logical and simple explanation for the source of the wing, but when dealing with the second par
t, namely, the evolution of the wing and of the muscular structure, the theory is rather general and presents certain problems if examined.
The alternate theory proposed in this essay can be considered as a modification of the paranotal theory in that while proposing a new source for the flight mechanism, one totally unrelated to flight, still, the paranotal extension is accepted as the wing so
urce. However, the alternate theory also generates possibilites for alternate sources for the wing, as well. For this reason the alternate theory is less simple and precise in describing the origin of the wing, but such lack of elegance is perhaps compensat
ed for by a rather simple origin for the flight mechanism and the development of aerodynamic lift and flight.
next para is an alternate wording of this para!
The alternate theory presented here is perhaps less elegant in defining the sources of the wing because it not only accepts the paranoia source, but even allows other possible sources for the wing. However, the proposed source of the flight mechanism, on t
he other hand, is simpler and more logical. The alternate theory, as will be seen, presents a perfectly acceptable idea for flight origins, which on examination appears perfectly obvious. It will be proposed that the mechanism and structure of flight derive
s from a source that has nothing to do with flight, and that flight is really an accidental by-product of a simpler, less design-dependant mechanism that was perfected before flight actually occurred. Such a development is in accordance with evolutionary te
ndencies, where developments in one system can lead to unexpected developments in a distantly related system. Examples of this are . . .
The paranotal theory, with its implied pre-adaptation, its straight, linear evolvement from primitively aerodynamic structures to an advanced aerodynamic function, flight itself, seems to have been created in hindview, as a backward un-developing of present
function and structure. This idea may not fit the unpredictable turns of evolution, and also presents serious problems when examined in detail.
To build an alternate theory, let us first look at several brief summations of the Paranotal Theory.
1. "The...widely accepted paranotal theory postulates that wings arose from lateral tergal expansions, or paranota, of the thorax. It is maintained that the prothoracic lobes of some of the most ancient fossil insects are organs of this kind which had pers
isted long after the paranotal of the other thoracic segments had developed into o wings. Paranotal expansions occur in positions characteristic of wings, not only in the thorax, but also on the abdomen in various arthropods; among insects they are seen in
many larvae and also in Lepisma, where they contain tracheae recalling those of a wing pad. It is suggested that they became sufficiently large to function as gliding planes or to control landing in leaping insects. Later they acquired basal articulations w
hich, along with the development of muscles, enabled them to become organs of independent flight. It is not improbable that at an early stage in their evolution wings were concerned more with sexual display or temperature regulation than flight."
There is also a tracheal gill theory, which is not accepted, because wings and gills do not share the same anatomical development and "do not seem to be serially homologous with tracheal gills". (R. G. Davies,
Outlines of Entomology, 1988 p. 24). It is also based on assuming that winged insects had aquatic ancestors, and this is not supported by the facts.
2. "It is now widely, but not universally accepted that insect wings arose from meso- and metathoracic paranotal lobes, homologous with those which occur on the prothorax of members of the Paleozoic order Palaedictyoptera..."
"In these ancestors it is supposed that the lobes later came to serve for delaying descent, as parachutes and then as aerofoils, those on the meso-and metathorax becoming enlarged and
MODIFIED UNTIL M WITH ASSOCIATED CHANGES IN THE SKELETON, MUSCULATURE AND NERVOUS SUPPLY OF THE THORAX, TRUE FLAPPING FLIGHT WAS EVOLVED. THE PROTHORACIC LOBES BECAME OBSOLETE and ULTIMATELY DISAPPEARED, AFTER PERSISTING FOR A WHILE IN A FEW LINES
." (R. J. Wooton, The fossil record and insect flight, in Rainey, p. 236)
"The paleontological evidence, slender as it is, seems to favour the existence in the Devonan a small -- c. 10mm? -- insects, whose initially protective
thoracic paranota became enlarged in association with their value first as parachutes, in delaying descent; next as gliding surfaces; then steering vanes, as they developed the ability to slightly pronate, supinate, el;evate and depress the pads by acti
on of the pleural leg muscles." (ibid. p. 248)
3. "At the present time there is almost universal support for the paranotal theory. Most authors suppose that the outgrowths were favoured in selection by their ability to serve as planes utilized in gliding from high vegetation to the ground. Hinton start
s further back and considers the first step towards flight to have been attitude control of legs and body to ensure that the falling insect landed on its feet and could make a quick escape from predators, namely spiders. Enlargement of lateral outgrowths wo
uld be a later step in making true gliding possible. Flower, who suggests that gliding could have begun in small insects of 1 cm length even before lateral planes appeared, points out that diminutive paranotal lobes at their earliest appearance could have p
layed an important role in attitude control.
"Then, as visualized also by Pringle, hinges to the planes and muscular control of their inclination would enable large adult insects to exercise control over the direction of gliding; and, finally, relatively slight alterations in the muscular system woul
d provide for slow flapping flight. That is probably the most widely accepted conception of the evolution of wings and flight. (V. B. Wigglesworth: The
Evolution of insect flight in Insect Flight, ed. R. C. Rainey, Symposia of the Roy Entom Soc of London, 1976).
4. The fossils from the Carboniferous indicate an abundant and probably rapidly evolving pterygote fauna, with species often of large size such as the Palaedictyoptera, dragonfly-like, with a wing span of up to a meter. There are abundant remains of wings
of many kinds and sizes, but there is no conclusive evidence as to where and how wings came into existence. It can be assumed that paired thoracic expansions first formed gliding organs. Some Palaeodictyoptera, such as Stenodictyon, possessed small prothora
cic dorso-lateral expansions as well as large mesothoracic and metathoracic wings, with wide bases, probably used for gliding. A narrowing of the wing base and added complexity in strusture could lead to the ability to fold the wings when not gliding, an ob
vious asset possessed by most winged insects today. Simple flapping flight presumably came after the ability to fold the wings... Wings develop\ ontogenetically as double layered outgrowths of the body wall on the mesothorax and the metathorax...Tiegs (1955
) pointed out that in acquiring the ability to fly the Pterygota must have used existing thoracic musculature of non-flying ancestors. (S. M. Manton,
The above quotes show a basis for flight is strictly built on jumping and gliding, that is, aerodynamic attitude control -- in fact, there seems to be no other way. However, by thinking in different directions we can arrive at a different set of conditions
that could lead to flight with far less complications.
EXAMINING THE PARANOTAL THEORY
The structural source of paranota.
Paratergal lobes refer to lateral expansions on any of the terga, or dorsal plates. Paranotal extensions, or paranota, refer to those occurring on the thoracic terga, called nota (singular notum).ILL
The presumed development of wings from paranotal lobes seems perfectly logical. The identification of this source rests on the fact that the anatomical structure for both is the same; that is, the two organs are homologous. Both are folds of the integument,
both are more or less extensions of dorsal surfaces, and both are located on the thorax. There are abdominal paranota as well, but this only underscore potential in arthropods to possess paraterga. Actually, the emergence of paranota, or more generally,
paratergal lobes, that is, lobes occurring on any tergum, presents no difficulties in its explanation: variation in size is a common biological occurrence, and paraterga are nothing more than increases in the lateral dimension of the terga. Unklike longitu
dinal tergal expansion, such extension need not meet any barrier because it expands away from the body. It is a fact that paratergal lobes exist widely among arthropods in general, and in both in fossil and recent insects. The basic plan
ILL 4 of a paranotal lobe is that of a folding out of the body cover at the lateral edge of the nota.
Paraterga are found on ILL 4 fossil Trilobita and Cheloniellida, and on Xiphosurida (fossil and recent horseshoe crabs). Paraterga may or may not be present on Crustacea. Fossil Arthropleurida (giant fossil myriapod has them. Among the Myriapoda, ce
ntipedes, pauropods do not have them, but most millipedes do carry recognizable paraterga. In the Insecta the lobes are mostly absent in Collembola, Diplura and Protura, but show up to a small degree, and the THysanura and Monura possess them on trunk segme
nts. The basic innateness of paraterga is evident not only in their frequent appearance, but in their repression and later derepression in some lines, as well. "Porthoracic paranota seem to have been redeveloped among some tropical mantids, and in the insec
t family Peloridiidae of the Homoptera. (Boudreaux, p. 136)
Being extensions of the sclerotized tergum, the paraterga are equally hard, therefore they effectively expand the protective area of the terga. Since they enlarge the horizontal diameter of the insect, they also increase apparent size and thereby would be
adaptive in defense, or competition.
At this point, therefore, a convincing structural source for the lobes can be demonstrated.
Size increase in paraterga
If not adaptively useful, novel structures tend to disappear. Paraterga, in general, can increase laterally to provide certain adaptive advantages. The paranotal theory has no problems here: a small series of minimal lateral extension of the nota readily i
ncrease the visual size of the insect. To an insect relatively small increments appear more significant than to creatures of our size. Therefore, it is possible for ancestral paratergals extensions to reach a size where they would have aerodynamic effects.
But if flight does not originate from jumping or gliding, then the paraterga would have no aerodynamic roles to play. In this case hinging would hardly occur because protection would not be maintained, although conceivably the insect would be able to exagge
rate its apparent size, yet still move less encumbered in narrow spaces.
We must distinguish between the primitive, sclerotized paraterga, and the apparently thin prothoracic paranota in the fossil record. The accepted theory speaks at first of an occurrence, a secondary development, of the general ancestral tendency to evolve p
araterga, and assumes them to eventually become thin lobes such as found in the Paleozoic Stenodictya and Lemmatophora. It has been said that "paranotal lobes are probably secondary and serve such functions as eliminating of shadow for camouflage, protecti
on against predators, or streamlining, any of which may also have been the initial function served by the primary lobes of Pterygote ancestry." (Wooton, p.236-237) However, this view does take into consideration the initial, primitive form of the structures
Camouflage is conceivable if the paraterga, whether ancestrally primitive or advanced to the lobe stage, are ILL5 longitudinally fused and laterally extensive, because than they would cover the legs to some degree, and make the insect resemble more a rand
om natural object, like a pebble, than a six-legged creature. But if the lobes were separated, not joined, as they appear on the reconstructed Paleozoic insects with prothoracic paranota, then such geometric spacing and form would actually
all attention to itself. The geometric designs on existing insects are the opposite of camouflage, they are visual identifiers.
Protection is an acceptable adaptive function of primitive paranota if they either visually enlarge or structurally protect. But protection implies density and mechanical strength, both of which are likely be absent in the lobes assigned to be gliding vane
s. Protection can be of various kinds, ranging from defense against larger predators to avoidance of physical abrasion.
We must also distinguish protection from predation from protection in interspecific contest. What offers little protection against larger predators, can be effective in rivalry among similarly sized conspecifics, where thin paranotal lobes, acceptable for g
liding, might provide advantage even if not particularly strong. The degree of protection from a complete range of sizes of aggressors decreases with the size of the insect to be protected by paraterga.
Streamlining must refer to aerodynamic surface control and would be effective once jumping and gliding were performed. If speaking of pre-glide streamlining the term must refer to ground motion. Here, again, sclerotized, down-curving lateral extensions of t
he terga would be acceptable, especially if closely fitting together on their anterior and posterior egads.
But here little gain would be made by a lightly structured, well-separated sets of lobes when moving through vegetation or narrow spaces.
Print from here
So far we have found the paranotal source of wings acceptable, and their initial, non-aerodynamic adaptive functions partly, but not totally plausible. Now we come to other, crucial developments for flight, such as jumping, parachuting and gliding, postulat
ed by the accepted theory as the fundamental, kinetic basis for flight..
Jumping in general
The assignation of jumping as the basis for gliding and then flight evolution at first the sounds perfectly reasonable. But let us consider some details, firstly the size factor. Except for the Orthoptera, existing jumping insects do not measure v
ery large. Collembola (springtail), Siphenoptera (fleas), Homoptera (spittle bugs (family Cercopidae), leaf hoppers (families Cicadellidae, Membracidae, Psyllidae), click beetles (Elateridae), cheese maggot (Larva of Piophila) are all minute to small insect
s. At their size any but the most haphazard gliding would be present practical problems because of the forces of aerial drag and air movement are large in relation to the insect's small mass with which to counter these. The simple fact that there are so few
jumping orders can be seen as indication of this problem. It is also significant that none of the jumping insects perform gliding.
Among the Orthoptera, in Caelifera, or grasshoppers, and on the Ensifera, or katydids and crickets, we find sizable insects that jump. Yet they -- who could easily do so with their highly controllable wings -- are not noted for gliding. Among all insects on
ly certain locusts, Lepidoptera and Odonata have been observed gliding, but this is clearly exceptional, and among these medium to large insects only the Orthoptera are jumpers. We can conclude from the existing evidence that gliding is not useful for insec
ts in general. Opening the wings as part of the an escape jump by grasshoppers serves not to glide, but to visually confuse the pursuer. And since the lobes were supposedly helpful in maintaining correct body orientation in jumping it is surprising that gra
sshoppers do not attempt any aerodynamic control: the fact is they are clumsy saltators, often landing upside down. They leap "...some rising in strong light, others tumbling aside with clumsy leaps." (Lanham, p.145). Other jumping insects pay little attent
ion to proper landing orientation; perhaps merely jumping with high enough speed is satisfactory not only for insects but also for frogs. The escape value of jumping arises from the fact that a pursuer's visual tracking ability is outclassed by the speed o
f the leaping insect: a movement of a sufficiently small object, travelling at sufficient speed, and occupying a sufficiently large angle of the observer's visual field can be impossible to track by a visual system.
NO VESTIGE OF EARLIER SALTATION IN OTHER PTERYGOTA!
An interesting question concerning jumping as precursor of flight can be asked. Why no patent or latent vestiges of jumping structure or behavior have been found in any form among the great many families of flying insects? Had jumping been part of the orig
inal shared ancestry we could reasonably expect it to have left traces or to have led to additional developments.
It can be remarked that among apterygotes, or insect orders without wings, jumping is the exceptional mode of escape or locomotion.
Jumping with general paraterga (not thoracic only)
Except for fleas (Siphanoptera), we think of insectan jumping as a form of escape behavior. But if jumping with lobes is presumed to lead to gliding and parachuting, we may point out that horizontal extension of body surface decreases the escape time somewh
ere in the trajectory.. Lobes would oppose some vector of the jump.. It seems that lobes offer little improvement in escape. It may be significant fact that no jumping insect has developed any glide control behavior or mechanism. Such absence may simply sig
nal that aerodynamic control in insectan jumping through evolutionary design is difficult, or not profitable to achieve. Fortunately, conclusions in this topic are easily available from experimentation.
--------------------- ALL THIS Critique of jumping and gliding NEEDS A LOT OF WORK STILL!!!!!!
There is also the question of the angle of the surface from which the jump is initiated, or the start-orientation of the jumping body. We can picture an hypothetical jumping, non-flying somewhat grasshopper-like insect with lobes to help it control attitud
e. ILLS HERE: jumping in different directions -ILL and text
1. Here we have a jumping insect with immovable lobes, starting with body parallel to the ground and maintaining it at the same attitude through the entire 45o trajectory. In the horizontal part of the trajectory the lobes will impede minimally, bu
t they would amply retard speed in both ascent and descent. At all three stages the lobes would of course stabilize the attitude -- but only if air movement was not unfavorable and the.
2. In a steep 70o, jump the forces opposing ascent and descent would be further increased and gliding or parachuting would by slowing the movement would present no advantage to escape..
3. In the case of a low angle jump the lobes be of minimally effective. They would mostly lie in an approximately horizontal plane and would help float the insect and therefore the escape speed would be still be decreased.
Moreover, concerning the body orientation at the start of the jump, it should be considered that insect may often be in a non-horizontal, even vertical positions on rocks or plants. Granted the vegetation in the days of pre-flight insects was different fro
m that of today, still, directions of plant growth most probably ranged in angles from horizontal to vertical, and so insects were likely to be found in a non-horizontal position. If we take a present day case of a grasshopper on a grass stem
ILL 9 we can see how horizontal paranotal lobes would be of little service to an insect either landing on a horizontal surface or on a vertical one.
The problem of pitching in jumping with thoracic paranota 1/2/1991
Because jumping requires relatively high speeds the paranota would meet greater air resistance, and since these lobes are situated on the anterior half of the bopdy, there woud be a tendency to cause pitching, or bodily rotation on the transverse axis.
(ILLS <<GT) Models can be made from 55 mm long wooden matchsticks, with igniting heads scraped off, with wings of varied sizes, made of paper and attached with tape, varying in weight from 250 mg. to 1500 mg (the latter being an acceptabl
e average locust weight). The sizes of the lobes range from those having minimal to maximal observable lifting effects. With smaller lobe-to-body proportions there is decreased pitch, but there is also decreased aerodynamic effect as well, so attention on l
arger lobes relative to the body size is permisssible when looking for aerodynamic effects.
At any rate, with such models we can demonstrate the problem of pitching. If we toss these models to reach various distances, up to 50 cm, launched at varying trajectory angles, we find that the front ends of the flying models have an almost constant tenden
cy to rotate up or down, causing the model to go into rotation and to crash.
Aside from the suggestion that in jumping, pitch rotation due to paranota may make their use counterproductive, there is an interesting point that emerges from these trials. If the models are turned around and are launched with the lateral expansions in the
posterior half, the pitching action is emphatically reduced, causing the model to complete its trip. If this observation is indeed applicable to jumping insects, then we can ask whether to improve attitude control in jumping, it might not have been appropr
iate for abdominal, rather than thoracic paraterga to develop. This question also arises in passive gliding (discussed below), therefore we may consider the point as unsupportive of deriving flight from paranota.
Running with lobes
Would lobes assist a running insect? We read that "<[>lobes] served as gliding planes and prolonged the motion through the air of insects able to run or jump." (Pringle, Flight/Locomotion, p. 434). Lobes would only assist a fast runner, but among present d
ay insects high speed runners tend to be ground dwellers, and their speed enables them to quickly reach hiding places.
Parachuting with lobes
THIS needs rehashing and cleaning up. cf GLDCHRT.SAM 1/2/1991
Parachuting, is an action that slows descent and so, in escaping, it may in some cases be useful, in others counterproductive. If the predator is large or mobile enough to still reach the insect that jumped off a height, for the insect to slow its decent wo
uld be a disadvantage. If, however, slowing the jump before reaching the targeted surface allowing attitude control in landing would be useful if the landing area provides escape (plants or crevices). Then, even if the predator is large and mobile, the inse
ct will gain an advantage in escaping. The effect or air movement is also influential. It may help increase the distance for escape, but it may may also negatively influence the parachuting by upsetting the attitude. And again, the weight and size of the in
sect must meet requirements.It seems that the adaptive value of parachuting is not clear-cut, and works only in a
narrowed set of circumstances.
NEW: BOUNCING!!
What would be the use It is in nuptial display that we can see a pragmatic use. Parachuting would be useful in a nuptial swarm, where a mass of male insects would, perhaps repeatedly, climb up on plants and would float down. Thu
s they would provided a visual signal to attract females since placing visual objects higher makes them visible from greater distance. Parachuting has also been assigned to a theoretical pterygote ancestor climbing up high plants to eat and then "descending
on the enlarge paratergal outgrowths. As Hocking notes, the appearance of winged insects and high vegetation was almost synchronous." (Sharov, p. 111) Such synchrony would indeed underwrite the relationship of the two. However, except for relatively short
plants, feeding on plants that may be considered high at least for insectan standards, there would be enough to eat both going up and coming down, thus there would be nothing gained in jumping off.
Gliding with lobes
THIs needs rehashing and scrapping! cf. GLIDECHART.SAM 1/2/1991
As for gliding in non-escape situations, (unrelated to aggression) there seems to be no convincing purpose. Insects generally have an interest in holding on to the substratum, while sudden dropping or jumping are escape reactions. In territorial dispersal t
here is advantage to gliding as long as there is enough air movement to carry the insect a significant distance. However such application puts size limitations on the insect and also requires that the glide be started from a significant elevation. Aerial di
spersal may not be a point general and crucial enough in adaptive success as more basic functions such as mate finding, mating attraction or territorial rivalry. There is a less than fully convincing argument in another basic proposition of the paranotal gl
iding theory, which speaks of insects climbing up and then gliding down to save energy. The practice of saving energy from having to climb down after having climbed up is not found in insect behavior. And the proposed initial emergence of high plants as a s
timulus for gliding is less than convincing for additional reasons discussed below
(under Support)<;>
Gliding, moreover is an uncommon practice among flying insects, except when there is a sufficiently large wing surface. "Occasionally insects are seen to glide with the wings outstretched, the lift forces produced by rising air being sufficient to keep the
m airborne without any wing movements." "Various Lepidoptera, locusts and Odonata have been seen to glide..." (Chapman, The Insects, 2nd ed., p. 196).
Just as parachuting, gliding would also have served a plausible purpose in nuptial behavior, where slowing the insect's movement through the air would cause its image to be easier to notice and to track. Still, this idea will remain just a seemingly obvious
fact, perhaps made irrelevant by actual experiments on exact aerodynamics involving the insect's size, the lobe's surface area, and the ballistics involved. Perhaps slowing down achieved by lobes below a certain size are insufficient significantly (in inse
ct perspective) to alter the speed of descent, in which case in the earliest part of its existence the lobe could not have been aerodynamical useful.
Once flight had begun, gliding would, of course, make perfect sense. At that time the expectedly primitive flight musculature would benefit from periods of rest during flight. The prothoracic paranotal lobes on the fossil
Stenodyctia may be representing a stage where relatively primitive flight control would need be assisted by additional gliding surfaces. <[>Perhaps prothoracic lobes developed only when needed to assist early flight reached through]
pbb out ! TO KOME: the idea that wings must be in frontal area, and if they were not, then prothoracic vanes would help this problem. ILL + text DO here: wing line: optic, etc.
Jumping must be initiated at a relatively high velocity for at least one reason: to carry the insect the maximum distance and so the pursuers' visual tracking becomes confused. But speeds proper for jumping might be too high for gliding all throughout a tra
jectory on the ground. When jumping out of higher vegetation there could be enough time for a small insect to slow down to a speed compatible with gliding, but it is uncertain whether the insect might still be travelling with head in front.
Why improve on jumping?
Even if a closely controlled and selected development of an aerodynamic surface to modulate jumping may be conceivable and available, the question remains: is there any reason to improve on jumping? Jumping is certainly an effective method of escape. An in
sect that leaps disappears because its speed is not successfully followed by the visual system of an observer, and as long as the insect is no longer seen, the jumping ability, whether ending with legs ready to grip a substrate or not, is positively selecte
d. After all, jumping is mostly rather successful for grasshoppers, at least from human attention. But if birds and mammalian carnivores are better at it, they were not around in the formative era of flight when on land only primitive vertebrates existed, a
nd predatory insects were equipped with nothing better than insect vision. If these conditions of predation prevailed in early times, jumping may have needed no improvement.
Looking at less advanced arthropods, we can find jumping millipedes in the genus
Dipsiulus, but they exhibit only a primitive form of jumping, performed on the ground, while running, reaching 2-3 cm, employing not so much the legs as an upsetting the center of gravity by curving the body upwards while in motion. Moreover, the mi
llipede peforms little control and "normally falls on its side." (Manton, Arthropoda, p. 336)
Among apterygotes, in the Thysanuran family Machilidae, the seashore bristletail,
Petrobius, performs escape jumps using its abdomen, not the legs, leaping "high off the ground, landing some 3-10 cm away...with the head pointing in any direction." (Manton, Arthropoda, p. 337)
The last two examples of jumping seem to indicate that such escape reaction is useful even when without any precision control over landing conditions.
Attitude control in jumping or gliding with lobes FLOWER! (this was redone on paper -- to be reconciled with this) 1/2/1991)
This needs work!!
Examining the literature, it seems that no experiments on jumping and gliding with relation to paranotal lobes have been conducted. The only study we can find is based on calculations involving aerodynamic air-flow affecting cylindrical objects dropped vert
ically. A horizontal vector, representing a 2 mph breeze added in order to produce a horizontal factor of gliding. (J. W. Flower, On the origin of fight in insects, J. Ins. Physiol, 1964, vol 10. pp. 81-88) The calculations predict that only an insect with
about 1 cm length would have any chance at acquiring gliding ability, Smaller or larger insects would find it less practical to maintain any control.
The optimal condition suggested is for a drop of 30* incidence, on a glide of about 45*, with an attitude placing the head 15* below the horizontal. The author states that cylindrical objects of optimal size would be in a range of aerodynamic factors where
a small alteration in body shape "such as the appearance of 'corners'" could "give moments large enough to effect the attitude of the insect..." (Flower, p. 87) and that such a shape-alteration could be provided by paranotal lobes. It is also concluded that
"attitude control of insects could be by the use of legs or by very small changes in body shape; the insects of around 1 cm long could have achieved a worth-while glide performance without wings, provided certain attitudes could be maintained; that control
by change of body shape is more efficient than by legs; and that from these first rudimentary wings larger wings would evolve, further improving the glide performance." (Flower, p. 88).
We must note that the above description of successful gliding with lobes depends on maintenance of suitable attitude. but this is where problems arise. Considering that attitude control has not been reached by any insect that jumps (with or without wings),
or glides (without wings), it is tempting to assume that such control is not feasible within insectan conditions.
In fact, a simple set of trials will show this. Models of insects with paranotal lobes can be constructed from paper and matchsticks and iron nails. GIVE DIMENSIONS AND WEIGHTS! After making a set of models of different weights (up to 1.5 gm -- roughly the
weight of an average locust), and of different amounts of lobe surfaces, it is only necessary to manually launch these at varying speeds and initial angles of attack. ILL<;>GT After conducting a large number of trials it becomes evident that the models cann
ot maintain attitude because they are subject to yawing, or rotation about the transverse axis of the body. ILL
There may be ideal, narrowly limited conditions where such rotation is minimal, but the such stringent limits posed would severely narrow the possibilities of glide development. This situation would not fit the ways of evolution, which most generally ap
pears to develop new structures in areas of open possibilities and wide tolerances.
These trials used bodies longer than 1 cm, but if we are to follow the findings of the above paper, a boy 1 cm long ought to be used. When attempting this it becomes obvious that at such size controlled ballistic conditions and filming is required to see th
e details of the action. When dropping or launching a 1 cm cylinder, weighing 53 mg, made of rolled paper, bound with masking tape, with or without appropriate lobes, the object and action is too small and too fast to follow, but a tendency to bounce when l
anding is clerly observable. We can drop or toss the model at various angles of incidence and in a rnge of heights from 10 cm to 50 cm, landing on surfaces of varying hardness. These can include a table top, a pad of note paper, i cm thick, the top leaf of
the pad curved concavely, or a (kleenex) tisssue paper loosely held horizontally in the air. There occurs a bounce on landing in almost all instances. the energy is only absorbed fully when the model falls vertically on tissue paper. Even tissue paper bounc
es the model when the incoming angle is not vertical.
The problem of bouncing recalls the clumsy landings of all jumping insects, and probably illustares that any success and advantage in jumping with lobes is confined to limits of rather narrow tolerance.
they ASSME A FLAT GROUBD FOR LANDING, ATTITUDE AND ALL THAT FOR QUICK ESCAPE. WH AB BUSDHES/
Problems in the derivation of flight from jumping and gliding
1. Existing jumping insect do not glide as a part of jumping. There seems to be little advantage gained. Grasshoppers might seem like an exception, however, their opening the wings briefly in mid-jump is primarily a visual defensive.
2. There are insects that glide, while in flight, in a manner not at all related to jumping. These insects (Orthoptera and Lepidoptera), have large wing surfaces, and such aerodynamic requirements are totally absent in relatively small surface areas assign
able to hypothetical paranotal lobes.
3. In existing insect paranotal lobes of any significance have not developed -- a fact not supportive of aerodynamic functional possibilities inherent in tergal extensions.
4. While the paranotal theory ascribes flight evolution to apterygote sources that glided, among existing apterygotes none glide.
5. There are examples of jumping among the more primitive apterygote insects, as in Machilida ((bristletails) and among the non-insectan millipedes, but these actions do not involve any functional gliding.
6. No apterygote jumpers have any paranotal lobes of aerodynamic significance.
7. An assumed ancestral jumping action has neither widely survived or further developed among pterygotes. The occurrences of jumping among Orthoptera, Siphanoptera, some Homoptera and the occasional Coleoptera provide no generalizable pattern to counter the
conclusion that in each of these insects jumping is a distinct, specialized and unrelated development.
8. The avenues of development are constrained and have small tolerances. Such conditions leave little room for possibilites for wide-ranging experimentation and variation, fundamentals in evolution.
The flight mechanism in brief
At this point let us briefly consider the flight mechanism. There are three types of muscles involved: direct, indirect and accessory.
ILLS 10 The wings are moved up by an elastic downward deformation of the thorax through the pull of the dorsoventral muscles connecting the sternum, or bottom and the dorsum, or top of the thorax. This is the indirect muscular mode. In the
direct mode, the wing is pulled down by the dorsopleural muscle which comes up from the sternum to the pleuron, or side, just under the wing hinge and through a short tendon it connects to underside of the wing. The small accessory muscles found at the win
g hinge area act serve to control the flexing the wing needed for directional control.
The dragonflies (Odonata) and cockroaches (Blattaria) fly the way just described. But there are two other variations. In moving the wings down the beetles (COleoptera) and grasshoppers, crickets and katydids (Orthoptera) add an indirectly acting muscle to t
he direct one. The indirect muscle, the dorso-longitudinal, contracts the thorax longitudinally and acts through a reversal of thoracic deformation caused by the other indirect muscles.
The third variation appears to show advanced development, evolved for high-rate wing beat on relatively small insects. The true flies (Diptera) and the wasps, ants and ? (Hymenoptera) move their wings both up and down indirectly through the alternating defo
rmations of the thorax. The direct muscles, attached to the base of the wings also act in twisting the wing as required for proper flight. Many of the direct wing muscles are attached to the coxa, or base of the leg and are in fact functional in leg movemen
The wing musculature system is complex, comprised of three different subsystems, and it seems somewhat removed from its purpose, as if it had been originally designed for another task. this implies that at as controlling the wings evolved, different muscles
were brought into use for different tasks. The indirect muscles are thoracic muscles which existed long before flight, and had basically played roles in skeletal muscular movement, used both in locomotion and in terrestrial breathing, and most of the direc
t muscles must have originally served only the legs. Nor would this fact be a problem since in evolution unexpected developments often occur -- such the descent of vertebrate jaws and hearing organs from gill arches are obvious examples. Nevertheless, it ma
y be said that in describing the evolution of moveable wings from stationary lobes there are no substantial clues offered by the paranotal theory to lead us to anything but the most general hypotheses.
Summary of the questions about the Paranotal Theory
The theory attempts to answer to the following:
1. The structural source of the wing
2. The adaptive role of the wing in its earliest appearance
3. The origin and development of the flight motor
The answer to the first problem is based on the homology between the existing wing and the presumed anatomy of the prothoracic paranota on Palaeozoic insect fossils. The presumption is reasonable, therefore we can accept the proposed origin.
The second topic has been questioned by us and some important points in the proposed functional development of the lobe and flight seem not as strong as would seem at first glance.
As for the flight motor, although this system is conceivably easy to generalize about (because it can be derived from earlier muscular functions, the questions raised, when examining the nature of jumping and gliding with lobes cause the seemingly obvious p
aranotal theory to permit a search for alternate theories.
LOOKING FOR AN ALTERNATE THEORY
A fresh view is sometimes hard to obtain when seemingly obvious answers to a question are no longer
satisfactory. We can perhaps start off with the paranotal lobe and try to find an alternate adaptive role that could have transformed it into the flying wing. In order to begin
from a relatively unbiased point, with no special direction in which to start, it might be helpful to stand back to take a general overview
NEW: We start off with the accepted ancestor of the wing, the paranotal lobe, but we will try to find (an) alternate adaptive role(s) for it. (Develop all this a bit longer)
In order to reach a relatively unbiased starting point, with no special direction in which to start off, it might be helpful to stand back and take an overview of a listing of insect behavioral functions.
Insect behavior can be classified and listed as:
Sensory perception
Basic responses and patterns of behavior (Reflexes, learned and instinctual behavior)
Periodicity and clocks
Orientation, navigating, homing
Host selection and feeding
Parental care and presocial behavior
Population behavior
Behavior, genetics, evolution and speciation
Eusocial behavior
Let us considered the accepted picture of the typical insect with proto-lobes used in jumping.
(ILL fossil proto-lobe ancestor (Lanham, eg).
How else could a such a paranotal extension of the tergum function adaptively? The paranota of
Stenodictya appear to be thin, winglike, although whether membranous or sclerotized is probably not decidable. Let us go back in time to where they were still primitive sclerotized extensions of the terga. What use could they have aside from protect
ive, and related topics mentioned above?
Examining communication behavior, we do find certain options, because we know of many small structures in existing insects that can play a role in sound production. We are talking about minute appendages, serrated, striated or otherwise designed for sound
production. It therefore stands to reason that a proto-lobe, being a small expansion of integument on the thoracic side can be rubbed by spines or pegs on the leg to produce a stridulatory sound. Doing so we, as a matter of fact could claim to have found an
oxypragma for the proto-lobe. For even at a minimal size, being only a surface irregularity, it immediately possessed a capacity toward adaptive applicability. The exact meaning of the term "immediate" is debatable, nevertheless, in the case of integument
al folding we can define it. Since the integument has a definite, measurable thickness, when folded it would be twice as thick, this being the minimal diameter of an evagination even as it first came into existence. As for the extension of its plane surface
, this would be determined by the extension area produced by a minimal folding.
ILL 11
With this point in mind, connecting the proto-lobe with sound production directs us to further examine the topic of insect sonification.
<[>write this better: The area of behavior where wings and their movement are directly involved in non-flight actions is communication. Wings take part in two varieties of communication, sound production and visual signalling. Let us tak
e each in turn in examining for possible useful leads. WHERE DOES THIS GO?]
Insect sonification
Sound production and associated behavior is widely distributed among insects. Functions of sonification include self-protection and escape, since many insects emit sound when captured or disturbed. Insects, such as crickets, advertise territorial claims by
sound. But the most frequently encountered and best known form of sonification is in courtship and mating display.
Importance of sonification
It is clear why the use of sound in commenting is important for insects. Their visual system is relatively limited and they are small, so to spot each other insects in a given environment would have to be relatively close. By sending and receiving sound wa
ves, they effectively increase their distance of communication. And because there is much variability in the ways sounds can be produced, this richness of variety in sonification, when used in mating communication becomes an important factor in creating ge
netic variability. Communicating by sound was probably a cause of quickened speciation for insects.
Insects have to see things at short distances, millimeters or centimeters away, and therefore they have a They cannot see far because a lens that has to see things at close distances FIGURE THIS OUT!!! WIDE ANGLE LENS VS TELEPHOTO
The wide spread use and variety of sonifying in insects
The significance of sonification is easily evidenced by the diversity of methods insects specialize in for sound making.
A great number of insects communicate with sound and they employ all available body parts to make sound.. Their discrimination between sounds and sound patterns can be quite refined. Mosquitoes can distinguish among different high frequency wing hums. some
grasshoppers, can synchronize their sonic emissions with that of nearby individuals.
Was sonification present in early, pre-flight insects? Flight and wings themselves are tools of visual communication, therefore pre-flight insects lacking these would have an even greater need for sound. Being permanently anchored to the substratum, that is
the ground or plants (or animals) would decrease the insect's visibility and size of area they could cover in their searches. For this reason it is can be assumed that the pre-flight world of insects was richly endowed with ways of sonifying. The fact that
a number of larvae and even pupae produce sound may indicate to what primitive stage sonification reaches back in the insect history.
If making sounds is therefore as wide-spread, perhaps(?) more so than flying, perhaps even more, if larvae and burrowing insects (Tenebroids??) use it, then it is likely that sonification is very ancient behavior and certainly PREDATE flying.
Various methods of sonifying
Stridulation (strigillation? - Pringle) is perhaps the best known method of insect sound making. Here scraping or rubbing together a body part against another with serrated surfaces produces a sound. Grasshoppers, crickets are outstanding stridulato
rs. The function exists in a great many Orders, and practically all body parts are employed. Stridulatory organs can be found on the head (Nitidulidae, Erotylidae). Some "tenebroids beetles (wood-boring?) have striations on the underside of the head." Mouth
parts often act as sonifiers: "Corixa americana rubs its front legs against the proboscis." There are clicking mechanisms on the underside of thorax, files under the prothorax rubbed by leg (Siagona, a scarabid). Ridges on the abdomen can be rubbed
by the edge of the wing cover (in some Scarabidae), or by the femur (Heteroceridae).
The legs are an important tool in stridulation. ("?)Some log-horned beetles (Prionus) rub hind legs along edges of the elytra. In a few aquatic insects legs are rubbed against the head Many long-horned beetles rub the rear edge of one segment of th
e thorax (usually the pronotum!) against another. Some Hymenoptera (ants, mutillid wasps) rub edges of adjacent abdominal segments against one another.
Table of sound-making organs and their combinations in insects
Source: B. Dumortier, Sound Emission Apparatus in Arthropods, in R. G. Busnell, ed. Acoustic Behavior of Animals
Even some larvae stridulate. Larval beetles, as in the Lucanidae, Passalidae and Geotrupidae, scrape the middle leg against a file surface on the third leg. In one species,
Passalus cornutus, what was once the third leg has now been reduced to not much more than a file.
ILL Stridulation is performed by the pupae of some Lepidoptera, or butterflies. Aside from Adventitious, sounds produced incidental to some body movement, such sonifying is found in ten families LEPIDOPTERAN LARVAE?? (Hesperiidae, Papilionidae, Lyma
ntriidae and Saturniidae, which rub ridges of one abdominal segment against another. The larva of the Epiophlebia (Odonata) rubs ridges on the abdomen by ridges on the hind femur.
Stridulation is found even among ants: Ponerinae, Dorylinae and primitive Myrmicinae use edges of the gaster and the petiole as a scraper and file set. Among true flies, or Diptera, the Tephritidae stridulates by vibrating the cubito-anal, or "armpit", sect
ion of the wing across rows of bristles on the third abdominal segment.
Another mechanism produces sound by vibrating a tymbal which is tight, elastic membrane. Cicadas create their loud buzzing through tymbalic vibration.
Tapping
Termite soldiers thump their heads against the ground. The death watch beetle
Anobium thumps its head on the side of its burrow. Certain grasshoppers tap their feet on the ground, and some cockroaches hit the ground with the tip of their abdomens.
Air piping
The mammalian source of sound, the modulation of air
stream is also used by insects for the same purpose. Of the few examples known are the May beetle and the bowfly which expel air through spiracles.
The wings as sonifiers
Vibration
And now we consider the wings in their sound making function. In communication the wings emit a humming o
r buzzing sound, either while flying or stationary, which their conspecifics recognize.
The Scarabidae, the Orthoptera, the Heteroptera, the Diptera and the Hymenoptera almost always produce a tone audible to humans. (Frost p. 201) Other vibrations may surpass the high r
ange limit of human hearing. /Wing vibration is known to be a sexual identifier, especially in mosquitoes
(DO!) I DIDN'T WRITE DOWN THESE EXAMPLES! /
Curiously enough sometimes the thorax is involved: a bumble bee with wings held will still emit a buzz by means of some vibration inside the thorax. One can presume that the wing muscles are vibrating. The piping sound produced by the
queen bees had been thought to be resulting from air forced through spiracles, but this "sound is probably produced by the vibration of the thoracic scl
erites (wenner, 1964) and so may be regarded as a vibrating membrane mechanism. (Chapman, The Insects,
1st ed!, p. 589)
In addition, Acilius sulcatus,
a water beetle, about 16-18 mm long, has been observed to generate sound as it warms up for flight, but without exhibiting any wing movement. (D. Leston,
J. W. S. Pringle and D. C. S. White, Muscular activity during preparation for flight in a beetle, J.
The wings
can also serve in stridulation. The common grasshopper sound, that of the short-horned grasshoppers, comes from rapidly moving the row of tiny pegs o
n the inner side of the hind femur against a hard ridge alongside the front wing. Others families(?) scrape wings together while hovering. Long-horn katydids produce
vibrations through an elastic membrane, the tympanum, part ??? the base of the right front wing. The crickets, of course, create their powerful sound as they rub together the special sound producing bases
of their front wings.
Paranotal lobes and stridulation
Since we are talking of wings with raspy surfaces, could we not reduce these wings to the size of the paranotal lobes, as on
Stenodictya and still have them perform something? Perhaps not, since they would extend out from the body and where it would be inconvenient for a leg to reach it. However, if the lobe was further decreased until it was only an irreg
ularity, a barely noticeable ridge on the surface of the thorax, then, we would find an analogue of elytro-femoral stridulation, where the proto-lobe, diminutive as it was, would be adaptively functional. The irregularities, spines on the legs would supply
the moving part of the stridulating set. The action might be combined with additional stridulatory areas on the abdomen. would immediately produce a new function.
But the minute proto-lobes could also enlarge to a degree. If the minimal proto-lobe developed into a long and narrow, and thin ledge, it could not only stridulate but amplify as well, as they do in modern katydids and crickets. Various ways of possible
for stridulation can be imagined. The immobile but adjacent lobes, could also be rubbed against each other, powered by intersegmental movement. And although this situation is not as likely as their employment as edges to be scraped by legs, such presumptio
n is in agreement with independently moveable wings, as in Odonata
Proto-wings directed backwards could present themselves as files to be rubbed by the femora better than one directed perpendicularly to the body. Such placement of early wings has been advocated by some authors, because the backward directed wing pads are t
he only ones found on fossil and recent nymphs. (Sharov, p. 110-111.)
Visual communication roles of wings
Today wings perform important visual roles. The colorful optical designs of Lepidopteran wings, especially when stationary serve a vital role in mate location, as well as in defensive mimicry. However, this function is also performed by a secondary wing dev
elopment, the closed elytra of beetles (Coleoptera), so that it is not so much the flight apparatus, but the exposure of a large visual area that is significant. The relatively slow mobility of the wings and the consequent visual transformations increase th
e effect.
Wings also increase the visual size of the insect, so that an individual will be easier to recognize, and a collection of insects such as a nuptial swarm will significantly increase its visually reflective surfaces due to wings.
Possible visual role of stridulating lobes
Going back to primitive lobes as stridulatory structures, it is plausible that they might increase in size through selection for modulating or amplifying the sound produced. But in this event at one point they would also become visual objects. And although
the stridulatory origin of lobes is the start, later, functioning as visual signals, the lobes could augment their size and develop mobility through selection for maximum communicating effect because a moving object attracts attention. The stridulatory fun
ction would be lost but an avenue towards flight would open.
We may even note in a general way that wings are still used today for visual and sound communication -- possibly an example of prehistory surviving, reminding us of the concept of birds as surviving dinosaurs.
Present day visual communications by wings
The a visual role assigned to the wings needs little documentation. Insect bodies and wings are often beautifully designed in shape and color and pattern for intraspecicif and interspecific communication.
The Odonata, whose fossils are thought to be the close to early flying insects, are a good example. In bright sunlight the large, reflective wings act like mirrors. Url Lanham writes: "In a spacious mountain canyon one may see the sun glinting on the wing
s of hundreds of dragonflies, from half a mile away, as two or three hundred feet high in the air, each patrols its bit of airspace." (The Insects, Columbia U Press, 1964, p. 185) Their positions at rest strongly support their visual use as well. At
rest true dragonflies, or Anisoptera, hold the wing straight out in a horizontal position, ILL, while the damselflies, or Zygoptera hold theirs vertical). It is reasonable to assume that territorial insects like the dragonflies would present their
wings as territorial signals and these positions optimal observation angles. ILL Moreover, the closing and opening of the wings is automatically a signal function of visual pulsation. Important visual clues are given among Odonata by various ways of bendi
ng the abdomen, and also by non-flight wing movements, but the wings must also serve in recognition at a distance either when flying or perching. The visual posturing of Odonata is underscored by the fact that males "resident in their territories usually di
vide their time between perching and flying. Orthetrum coerulescenc flies for about 19% of the time," (P. L. Miller, Dragonflies of North America, 1987, p. ??) which means they perch exhibiting their wings 81% of the time.
The highly specialized, oversized eyes of the Odonata support our recognition of their extensive use of visual data.
The wings are highly functional visual signallers in other Orders. For example, Drosophila males exhibit mating displays composed of body orientation, tapping, wing vibration, wing flicking, wing waving, wing semaphoring, wing scissoring, leg vibration, l
eg rubbing, circling, licking, and finally mounting. (H. T. Spieth, Courtship behavior in
Drosophila, Ann. Rev. Entomol., 19: 385-405, 1974; quoted by M.D. Atkins in
Introduction to Insect Behavior, Macmillan, 1980, p. 112)
Evidently wings possess effective visual function and no doubt wings, offering both large visual areas and reflective surfaces, whether movable or not must have possessed a meaningful role in insect behavior preceding flight. The functions of light refl
ection, or dynamic wing motions are subject to selection and thus the wings will tend to develops selectively for structural strength and size of surface area.
Stridulating lobes with additional visual roles
At this point we can evaluate the hypothesis of the proto-lobe which through sonic use could also become a visual communication organ. Once the lobes have reached a visual applicability a mechanism to move them would greatly enhance their effectiveness.
As soon as the proto-lobe or lobe reached a certain size and so entered the limits of insectan visual perception, it could, especially, if moving, function as an effective visual signaller. In this role the lobe would be selected in a competition for increa
sed visual output.
Critique of the stridulating lobe
This theoretical development of a paranotal lobe from stridulatory surface actually supports the accepted paranotal theory. Lobes could indeed have come into existence as originally minimal extensions of the integument. They could easily strigillate when sp
ikes on the leg were drawn against their plain, stiff edges, but they could also have gained rugose or serrated edges. Perhaps some section of the stridulatory expansions could have evolved into resonating plates. Or still being immobile expansions but loca
ted on adjacent body segments the emerging lobes could also rub against each other through segmental movement. This could conceivably even evolve into the flight mechanism. ILL However, these propositions are dependent on a new set of speculatory condition
s, and they increase, rather than diminish the number of unknown factors we have started with. it may be is advisable to look for simpler modes of evolution.
PUT ELSEWHERE:
Perhaps before the existence of the vibrating thorax and the lobe some insects had visual patterns, colored patches or bright shining surface on the thorax. When sound producing structures appeared, these spots might pulsate, move slightly in the visual fie
ld. There will be more top support this idea later.
5.17 In effect, the sound production and the visual signalling developed simultaneously, in close correlation within the limits defined by their share of developmental requirements. Each one benefitted its output if the lobe surface area or surface reflecti
vity/smoothness increased. But the visual function tended to decrease and so was in some part selected out in favor of sound or flight<;>
5.19 As for the argument that the phylogenetically primitive fliers, the Odonata have no click mechanism (MS p. 11) THEY most likely DO! Orthoptera certainly do have it! see:...? They are probably not primitive either!
end of Flight _A
Basic motion of the insect wing in insect with an indirect flight mechanism Sheme of dorsoventral cut through a thorax segment with wings
a wings
b joints
c dorsoventral muscles
d longitudinal muscles
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