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Re: Fw: Dinosaurs and birds



Note: I first began thinking about the evolution of flight about 1965, and my 
ideas jelled about 1995, and have changed little since. I see 2 major problems 
specific to ground-up flight scenarios: 1) the establishment of a phenotype 
that allows synchronous flapping of the forelimbs while simultaneously using 
the hind limbs in alternating fashion. 2) crossing the threshold between zero 
and minimally beneficial forelimb aerodynamic locomotive assistance. The 
comments below assume those magical thresholds have been previously breached.

Also I try not to use the term "wing-assisted" because I feel the need to 
remember that the process discussed _culminates_ in a wing... so fore-limb 
assisted is the term I use, and is synonymous here w/ "wing-assisted". Comments 
etc., are numbered for clarity.

Don

From: Michael Habib <mhabib5@jhmi.edu>
To: dinosaur@usc.edu
Sent: Thursday, April 5, 2007 11:22:11 PM
Subject: Re: Fw: Dinosaurs and birds

On Thursday, April 5, 2007, at 07:12  PM, don ohmes wrote:

> I think I understand the "they-can't-run-any-faster" point/analogy 
> quite well. The analogy isn't apt, nor are your assumptions. You 
> assume that an animal that can generate forward thrust directs it 
> exactly forward/parallel to the ground, and further, cannot quickly 
> re-direct that thrust.

I don't actually make those assumptions.  Those are features for the 
model in which thrust from the wings is used to increase maximum speed, 
and I included them to show that such a limited model is not feasible. 

============================
1). Huh? 'Cant-go-any-faster-than-the-hind-legs-can-run" is the model/analogy 
_you_ use to "refute" the idea that forelimb assistance can convey advantage by 
increasing maximum speed. The assumptions I listed are inherent to _your_ 
argument that fore-limb assistance can't increase maximum speed in the absence 
of incline. They are _not_ features of the model I use for assessing the 
possibility of advantage gained by increase of maximum speed through use of the 
forelimbs. The model I use is qualitative and involves multiple cases for 
hindlimb traction, incline, and density of the locomotive environment (both 
atmospheric and vegetative). In the case of vegetation, one must consider the 
vertical gradient; it is very different to walk through waist-deep grass as 
opposed to ankle-deep, for instance. Perhaps you meant to say something else. 

In any case: (another) major flaw in the model that is used to 'prove' that 
inclines are required for fore-limb assist to be useful in forward locomotion 
is yet another underlying assumption; that of hard smooth ground. An animal 
that can create fore-limb thrust is _highly _ unlikely to
restrict itself to a theoretical parallel-to-earth-surface direction
for that thrust, and directing thrust slightly upward can increase
stride length and height w/out necessarily decreasing stride frequency. 
Conditions that reduce hindlimb traction even slightly such as muddy ground, 
shallow water, and certain types of vegetative cover therefore alter the 
benefit profile of fore-limb thrust considerably. I call this the 
'tread-lightly factor'... and it is easy to construct scenarios where _maximum 
forward speed_ is increased through _fore-limb assistance_. "Sinking in is not 
a nimble thing to do" -- a song someone should have written, but didn't.

A model wherein the simplifying assumptions are hard, smooth, flat ground and a 
constant thrust vector at precise right angle to the gravitational vector is 
interesting as a first step, and indeed inevitably results in a faceful of dirt 
or reduction of velocity for the poor creature required to operate under those 
conditions. However, when the results are applied to the real world where other 
conditions exist, it is a clear-cut case of garbage in, garbage out. 
============================

By contrast, I quite agree that generating lift forces in other 
directions, or in short pulses, might have advantages for agility or 
maneuverability.

=============================

2. "Might"?
============================== 



> You also assume that the thrust generated is constant in force, 
> another major flaw of the car analogy. You further assume that for a 
> speed increase from fore-limb assistance to be advantageous, it must 
> occur at full running speed.

If it doesn't occur at full running speed, then it still isn't helping 
from a total velocity standpoint.  

==================================
3). So? We are discussing a selection-driven process that turns forelimbs into 
forward flapping wings. An advantage is an advantage. Acceleration counts, 
particularly in combination w/ advantageous change in vertical position.
==================================

If the animal wants to run faster, 
it is more efficient to speed up the hind limbs than to add thrust.  

==================================
4). That depends _entirely_ on the ambient environment, bodyplan and lifestyle 
of the animal in question. Again, there is the implied assumption is that the 
process of evolving flapping forward flight through forelimb-assistance begins 
w/ some sort of bipedal cheetah analog running on hard smooth ground. Animals 
in the process of optimizing the hindlimbs for speed have 'chosen' an 
evolutionary path that is unlikely to result in flapping flight. Animals that 
need to approach a prey item from a slightly elevated position in 
less-than-perfect traction (for example) have an entirely different take on 
"efficiency".
=================================

Thus, regardless of the gait, aerodynamic thrust is not helpful for 
faster speed.  

=====================================================
5). Again, that depends on the capabilities of the hind limbs, the direction of 
thrust, and the physical qualities of the substrate. If by 'chance,' some 
benefit occurs, evolution toward flight continues.
=====================================================

It can be useful for increasing _acceleration_, and thus 
getting to a given speed more rapidly.  

=====================================================
6). Which, btw, is "a cursorial mechanism by which forward progress is directly 
enhanced by wing oscillation", and significant from the standpoint of a 
selective process.
=====================================================

Though, the efficiency of this 
technique is going to depend on the speed regime, planform, and a few 
other factors.  Some ground birds do use wings to burst accelerate 
occasionally, especially if startled.  Whether or not the force is 
produced in pulses or is constant is not particular important to the 
model.  For maneuvers (including fast starts), a pulse of near-constant 
force is usually going to be most efficient.

> None of the above assumptions, although necessary for mathematical 
> analysis, are correct in an _evolutionary_ context.

They are not required for mathematical analysis, actually, and I did 
not mean to imply such assumptions.  

==================================
7). Huh? If you are using a model, you have to take responsibility for the 
underlying assumptions.
=================================

Is there a reason that you 
separate mathematical/mechanical analysis from evolutionary analysis?  
I am generally used to melding the two together.

===============================================
8). You just have to pay attention to which form of analysis is subjugated to 
which. For instance, I think you sometimes forget that the relative efficiency 
between competing iterations of a given system at a given time is what is 
relevant in the competitive context, as opposed to the theoretical efficiency 
of the total system relative to the ambient environment. 

_For instance, consider this quote: "If the animal wants to run faster, it is 
more efficient to speed up the hind limbs than to add thrust." -- Mike H._

I agree that is a _generally_ true statement relative to the physics and theory 
of locomotion, but it is not necessarily relevant to selective regimes on 
specific bodyplans, or in specific environments.

===============================================

> Thrust can be, and is, used to increase stride length (including, but 
> not necessarily at, maximum stride frequency), decrease stride length,

Using aerodynamic thrust to increase stride length is probably not 
particularly helpful, especially at the maximum for the hind limbs, 
unless you mean to imply that the animal is actually leaping.

=================================================
9). "Particularly helpful" implies that you are capable of quantifying 
selective advantage, and determining a threshold of evolutionary significance 
for same, and I don't think you or anyone else can currently do that. Advantage 
is binary relative to construction of a specific evolutionary scenario, ie, it 
is "helpful", or it is "not helpful". If you find that it is helpful, it must 
be included in consideration of the effects of selection on heritable 
morphological variance. If you say "not helpful to increase stride length", 
then it seems to me inescapable that you are using
the unrealistically limiting assumptions: 1)  thrust is always at exactly right 
angle to
the gravitational force, and 2) substrate conditions are optimal for hindlimb 
traction. 

And why would I leave out leaping? If it works, it works. Prey capture 
scenarios come to mind, as do refugia.
================================================

> overcome obstacles, increase/reduce velocity and seek refuge (or more 
> generally, improve tactical position). These exploits convey 
> advantage, and there is no such thing as an insignificant advantage, > or
>  "narrow margin" in the evolutionary context.

Overcoming obstacles is a very reasonable use of incipient flight 
abilities, as is seeking refuge.  Changing velocity (ie. acceleration) 
is also quite helpful, though increasing velocity past what the hind 
limbs can produce is less mechanically feasible.

=====================================================
10). "Less (...) feasible" strikes me as being in the same category as 
"particularly helpful". And again I ask, "What is the traction?"
=====================================================

> Analogies drawn from adult modern birds are not very useful in 
> constructing quadruped-to-neornithine evolutionary scenarios, in my 
> opinion.

I agree.

> Juvenile quail and turkeys are another story, and observations of 
> wing-assisted ('fore-limb assisted' in the evolutionary context) 
> locomotion on flat ground are easily reproduced, and include all of 
> the exploits listed above.

We must be careful, however, because even juvenile galliforms are not 
particularly good models.  

================================
11). But probably the best living models we have, yes?
================================

They have a very advanced upstroke system 
(even compared to other living birds), and fairly hefty pectoralis 
mass.  

===============================================
12). The wing kinematics changes have been studied through the entire 
maturation process? Or are you extrapolating from adult birds? This comment 
(unlike other sections of this post) is not in any way adversarial... I am 
genuinely curious, and your expertise is impressive, to say the least.
===============================================

But yes, they do use the wings briefly for some maneuvers on the 
ground.  These should be kept in mind during evolutionary analysis, but 
we must also be aware of what mechanical limitations exist for basal 
birds and near-avians.  Juvenile galliforms have many derived 
characters not seen in basal forms.

> There is an  'assistance gradient' as the wings develop that ranges 
> from zero contribution from the fore-limbs to full flight, and a 
> narrow time window (in the early stages) involves very high wing 
> loading and (I assume) forward impetus ("thrust") only. The period 
> from 'zero contribution' to 'thrust only' is relevant to evolutionary 
> scenarios.

Of course; I agree with all of the above.  The question then becomes 
which sorts of assistance are mechanically feasible and which are not.  
Increasing maximum running speed happens to be a low feasibility 
dynamic.  

=============================================
13). In my opinion, that ("feasibilty" of increasing max run speed) cannot be 
determined by theoretical analysis; once the possible/impossible threshold is 
breached, subject to reasonable assumptions, it is time to take data.
=============================================

By contrast, changing acceleration, improving turning radius, 
or performing incline runs are all mechanically feasible ways of using 
incipient wings, though how helpful they are depends largely on the 
flight apparatus in place.  With an understanding of the flight 
apparatus and structural strength of basal birds, we can narrow the 
range of possibilities to further evolutionary analysis.

>  I have observed chicks in the wild, and the ones that flap get 
> further down the road than the ones that don't, especially quail. That 
> is anecdotal, but I still have some money, if y'all are betting men...

I wouldn't be surprised; they're probably using their wings to get to 
top running speed more quickly, or aiding balance (or both).  I have 
observed a rather wide range of birds, juvenile and adult, in both the 
wild and captive situations.  I have seen precocial chicks use wings 
for acceleration when startled from standing still.  I have not seen 
any evidence that they reach higher speed.  I'm willing to bet that 
maximum speed is not increased in the quail you observed, either.  

=================================================
14). So your opinion is that within a cohort of same-age quail, those whose 
feathers are continually pruned will continue to arrive at point B at the same 
time as those who have begun to receive thrust assistance from decreasing 
'limb-load'? Obviously, divergence will occur. But you feel it will not be 
measurable before the un-pruned birds attain flight?
=================================================

Nonetheless, better acceleration is helpful (which is probably what 
they are gaining), and another reasonable trait for selection of 
flapping.  Again, I only was referring to the hypothesis of increased 
maximum speed previously, not any other advantages of wings on the 
ground, of which there are a few.

> You may think 'fore-limb-assisted' scenarios for evolving flapping 
> flight that include inclines are more convincing, more probable, or 
> more efficient, and I would agree.

I'm not sure if they are more convincing or not.  I am actually 
somewhat critical of the WAIR hypothesis, as I mentioned in my original 
post.  Inclines are required for the specific WAIR dynamic, but not for 
other advantages of incipient wings.

>  For the reasons given in the 4th paragraph, I say inclines are not 
> necessary. And that most definitely includes scenarios in "which 
> forward progress is directly enhanced by wing oscillation".

What are you suggesting they are "not necessary" for?  

============================================
15).  I have not "suggested" anything. I have clearly and unequivocally stated, 
from the get-go, inclines are not necessary for valid fore-limb assisted (= 
"wing-assisted"), ground-up evolutionary scenarios. I paste in the original 
statement and your original response for those who might be confused about what 
started this:

I wrote in reply to one of Scott's posts: "Not sure I understand, from the 
perspective of a 'ground-up' selective
process that can transform a terrestrial mud-lover into a barn swallow,
where the line between volancy and various forms of wing-assisted
running is (inclines are NOT necessary, in my opinion)."

Mike H. replied-- "The incline is necessary, because without it there is no 
requirement to 
produce a lift force towards the substrate, which is the critical 
aspect of wing assisted running." 

My position-- Inclines are not necessary for constructing valid ground-up 
scenarios wherein forward flapping flight evolves; that includes, but is not 
limited to, scenarios involving selective advantage conveyed by forelimb 
generated thrust that increases the maximum speed of the animal.
============================================

If you mean 
maneuvers, clearing obstacles, burst accelerations, or balance, then I 
agree.  But those do not usually _directly_ enhance forward progress, 
with the exception of burst acceleration forward.  I should have 
included that exception in my original statement.  That said, my point 
was that inclines are important to WAIR dynamics

=================================
---- Yes, they are.
=================================

and that maximum 
speed can rarely be enhanced by wings.  

====================================
--- Sigh. "Rarely"?
===================================

Those are the two kinematics I 
had in mind with regards to forward progress being directly enhanced, 
which is why I said inclines were important.  I did not mean to imply 
that inclines are required for the evolution of flight from a cursorial 
ancestor at any general level.

> (If, and only if, "wing oscillation" is what we called 'flapping' down 
> on the farm. If not, what the heck does it mean?)

Yes, it means flapping, essentially.  I said wing oscillation to be a 
bit more broad, since flapping usually means a true flight stroke 
(though it doesn't actually have to, it does have that connotation).

Cheers,

--Mike H.