330 Mya Eurypterid trackway
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Dunk - 30 Nov 2005 19:30 GMT http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html
Nature 438, 576 (1 December 2005) | doi:10.1038/438576a Palaeoecology: A gigantic fossil arthropod trackway
Martin A. Whyte
A unique, complex trackway has been discovered in Scotland: it was made roughly 330 million years ago by a huge, six-legged water scorpion that was about 1.6 m long and a metre wide. To my knowledge, this is not only the largest terrestrial trackway of a walking arthropod to be found so far, but is also the first record of locomotion on land for a species of Hibbertopterus (Eurypterida). This evidence of lumbering movement indicates that these giant arthropods, now extinct, could survive out of water at a time when the earliest tetrapods were making their transition to the land.
==== dunk
Don Kenney - 30 Nov 2005 22:52 GMT >http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html > [quoted text clipped - 15 lines] >==== >dunk On a somewht related subject, Scientific American currently has a nice article available online on the evolution of tetrapods http://www.scientificamerican.com/article.cfm?chanID=sa006&colID=1&articleID=000 DC8B8-EA15-137C-AA1583414B7F0000
John Brock - 01 Dec 2005 01:51 GMT >http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html > [quoted text clipped - 12 lines] >now extinct, could survive out of water at a time when the earliest >tetrapods were making their transition to the land. Hmmm.... In a way this sounds like an arthropod analog of one of those big flat sprawling early amphibians. It makes me wonder what the arthropods might have evolved into if the tetrapods had never come into existence. Do you think they might have gotten really big? Yeah, I've heard all the reasons why arthropods aren't supposed to be able to get big. But the tetrapods needed many innovations in order to adapt to dry land -- without the tetrapods keeping them down, might not the arthropods come up with a set of innovations of their own?
 Signature John Brock jbrock@panix.com
John Harshman - 01 Dec 2005 02:00 GMT >>http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html >> [quoted text clipped - 18 lines] > come into existence. Do you think they might have gotten really > big? Define "really big". How about the 8-foot-long Devonian millipede?
> Yeah, I've heard all the reasons why arthropods aren't supposed > to be able to get big. But the tetrapods needed many innovations > in order to adapt to dry land -- without the tetrapods keeping them > down, might not the arthropods come up with a set of innovations > of their own? John Brock - 01 Dec 2005 02:24 GMT >>>http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html >>> [quoted text clipped - 12 lines] >>>now extinct, could survive out of water at a time when the earliest >>>tetrapods were making their transition to the land.
>> Hmmm.... In a way this sounds like an arthropod analog of one of >> those big flat sprawling early amphibians. It makes me wonder what >> the arthropods might have evolved into if the tetrapods had never >> come into existence. Do you think they might have gotten really >> big?
>Define "really big". How about the 8-foot-long Devonian millipede? I'm thinking more like cow.
Or brontosaurus. :-)
Or Mothra. :-) :-) :-)
>> Yeah, I've heard all the reasons why arthropods aren't supposed >> to be able to get big. But the tetrapods needed many innovations >> in order to adapt to dry land -- without the tetrapods keeping them >> down, might not the arthropods come up with a set of innovations >> of their own?  Signature John Brock jbrock@panix.com
Nic - 01 Dec 2005 02:31 GMT > >>>http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html > >>> [quoted text clipped - 22 lines] > > I'm thinking more like cow. I see - something more the size of a termite mound.
> Or brontosaurus. :-) > [quoted text clipped - 5 lines] > >> down, might not the arthropods come up with a set of innovations > >> of their own? anon1@sci.sci - 04 Dec 2005 05:40 GMT > something more the size of a termite mound. That gives me an iead. You all know that children's game "freeze" where one person faces away from all the others trying to sneak closer? Whenever the one person suddenly turns around, everyone must freeze. If any of them are seen moving, they must go back to their original start.
Now think of a mobile termite mound as a sort of Trojan horse. When the victim isn't watching, the termite mound creeps forward. But every time the victim looks at the mound, it's sitting there motionless, so the victim isn't aware of the attack in progress. Then when the termite mound is right upon the victim, the hoard of termites sneak out to attack. Various scenerios: (1) They all attack at once, overwealming the victim faster than it can run away, like army ants attacking. (2) Only a few termites attack at a time, and the victim isn't aware they came from the termite mound, so doesn't move further away from it, allowing additional attacks until the total amount of toxin is enough to disable the victim. .
Don Kenney - 03 Dec 2005 14:10 GMT >>http://www.nature.com/nature/journal/v438/n7068/abs/438576a.html >> [quoted text clipped - 17 lines] >down, might not the arthropods come up with a set of innovations >of their own? I think that conventional wisdom is that two factors keep terrestrial arthropods smaller than Moose, Elephants and saurians.
1. Arthropods have exoskelatons rather than endoskelatons. That's, I'm told, not an optimal way to build large structural elements like legs and grasping appendages which are subject to stress in a lot of directions; inadvertant blows from branches and such at odd angles, etc. I can't confirm this. I'm not a structural engineer and also it seems to me that engineers are constantly coming up with new ways to build light strong structures that differ from past conventional wisdom.
2. There is clearly a problem with getting Oxygen into a large animal and distributed all parts of a massive body. Vertebrates have solved this with a complex system of pumps, pipes, (where appropriate) check valves and compact gas-fluid exchange "devices" with very large effective surface areas. I have no idea if the arthropod "breathing" mechanism can be scaled to the needs of a very large animal. My impression is that it's not all that obvious that it can be done.
A third question is how a massive arthropod would dump metabolic heat. This is a problem for terrestrial vertebrates and might be more of a problem for a large critter with an exoskelaton massive enough to support the animal.
Personally, I'm not happy with digging up live arthropods even 10 cm long. They generally appear to be displeased with being disturbed, and entirely too many of them are equipped for retaliation. I'm just as happy that arthropods that size are rare and that most of them don't get much bigger than that.
Stanley Friesen - 03 Dec 2005 15:03 GMT >I think that conventional wisdom is that two factors keep terrestrial >arthropods smaller than Moose, Elephants and saurians. [quoted text clipped - 7 lines] >build light strong structures that differ from past conventional >wisdom. I, too, am somewhat skeptical of this.
>2. There is clearly a problem with getting Oxygen into a large animal >and distributed all parts of a massive body. Vertebrates have solved [quoted text clipped - 3 lines] >mechanism can be scaled to the needs of a very large animal. My >impression is that it's not all that obvious that it can be done. There is not just *one* arthropod breathing mechanism. The most widespread is the one found in insects. That one almost certainly *cannot* be scaled up. However arachnids have a different mechanism, called book lungs. I suspect this mechanism might prove more scalable.
>A third question is how a massive arthropod would dump metabolic heat. >This is a problem for terrestrial vertebrates and might be more of a [quoted text clipped - 6 lines] >as happy that arthropods that size are rare and that most of them >don't get much bigger than that.
 Signature The peace of God be with you.
Stanley Friesen
anon1@sci.sci - 04 Dec 2005 06:16 GMT > However arachnids have a different mechanism, called book lungs. > I suspect this mechanism might prove more scalable. So those giant spiders in King Kong could be real, on some unexplored island, such as Flores? :-) And maybe during the next bird-migration season, a baby giant-spider will hitch a ride on a bird and end up in Alaska, and switch birds to make it to the USA, and land somewhere away from humans where it can eat everything it can find for a few months until next Summer it's adult size, where it'll hitch a ride on the back of a truck and pop off the truck in MY TOWN, just down the street from MY HOUSE, and in a few minutes it'll be at MY DOOR, where I'll go check what's making the funny scratching noise, and when I open the door IT'LL EAT ME, and suddenly the posts from anon1 will be all GIANT SPIDER POSTS!!!
Or maybe it already hitched a ride last season, and already got to Alaska in July, and already got to the USA in August, and already grew to full size in late November, and already got to ANON1'S TOWN a half hour ago, and already came to ANON1'S DOOR ten minutes ago, and already ATE ANON1 nine minutes ago, and I AM THE GIANT SPIDER TYPING THIS RIGHT NOW, and I want everyone seeing this to tell me where you live so I can come eat you too!!!!! .
Mark Isaak - 03 Dec 2005 22:22 GMT >I think that conventional wisdom is that two factors keep terrestrial >arthropods smaller than Moose, Elephants and saurians. I think what you call "conventional wisdom" is used to explain why the giant ants and spiders in 1950's horror movies could not really live.
>1. Arthropods have exoskelatons rather than endoskelatons. That's, >I'm told, not an optimal way to build large structural elements like [quoted text clipped - 4 lines] >build light strong structures that differ from past conventional >wisdom. I strongly doubt this explanation. Some insects already have bits of exoskeleton that extend inwards, for things such as muscle attachment. Making more of them should not be a problem for evolution.
>2. There is clearly a problem with getting Oxygen into a large animal >and distributed all parts of a massive body. Vertebrates have solved [quoted text clipped - 3 lines] >mechanism can be scaled to the needs of a very large animal. My >impression is that it's not all that obvious that it can be done. Spiders have a complex "book lung" organ, too. Again, I don't see oxygen stopping evolution of giant arthropods.
>A third question is how a massive arthropod would dump metabolic heat. >This is a problem for terrestrial vertebrates and might be more of a >problem for a large critter with an exoskelaton massive enough to >support the animal. Again, I suspect the mechanisms used by mammals and reptiles could be adapted for arthropods if they really needed it.
I suspect the main factor limiting arthropod growth is the actual growth part. For an arthropod to grow bigger, it has to discard one exoskeleton and then grow a new one. That can get expensive if you have to do it twenty or thirty times to reach an appreciable size. And then you have to spend at least some time with little or no skeleton, which does not sound like pleasant state to be in, even for a few minutes.
-- Mark Isaak eciton (at) earthlink (dot) net "Voice or no voice, the people can always be brought to the bidding of the leaders. That is easy. All you have to do is tell them they are being attacked, and denounce the pacifists for lack of patriotism and exposing the country to danger." -- Hermann Goering
John Brock - 04 Dec 2005 01:44 GMT >>I think that conventional wisdom is that two factors keep terrestrial >>arthropods smaller than Moose, Elephants and saurians.
>I think what you call "conventional wisdom" is used to explain why the >giant ants and spiders in 1950's horror movies could not really live. Yes, elephant sized ants that are still proportioned like ants are as impossible as elephant sized mammals proportioned like ants!
>>1. Arthropods have exoskelatons rather than endoskelatons. That's, >>I'm told, not an optimal way to build large structural elements like [quoted text clipped - 4 lines] >>build light strong structures that differ from past conventional >>wisdom.
>I strongly doubt this explanation. Some insects already have bits of >exoskeleton that extend inwards, for things such as muscle attachment. >Making more of them should not be a problem for evolution. I was thinking along the same lines. Perhaps they would have acquired some mix of internal and external support.
BTW, when bone first evolved in chordates, wasn't it originally as an exoskeleton? Didn't internal bones come later?
>>2. There is clearly a problem with getting Oxygen into a large animal >>and distributed all parts of a massive body. Vertebrates have solved [quoted text clipped - 11 lines] >>problem for a large critter with an exoskelaton massive enough to >>support the animal.
>Again, I suspect the mechanisms used by mammals and reptiles could be >adapted for arthropods if they really needed it. [quoted text clipped - 6 lines] >skeleton, which does not sound like pleasant state to be in, even for >a few minutes. Clearly a cow sized arthropod couldn't molt! But couldn't the external skeleton just grow, the same way internal skeletons do?
The interesting thing to ponder though is that *something* would have happened if arthropods had had the land to themselves, and that it would probably have included arthropods *very* different from any that every actually existed. Looking at Eryops, who could have foreseen a monkey or an eagle?
 Signature John Brock jbrock@panix.com
r norman - 04 Dec 2005 02:25 GMT >>>I think that conventional wisdom is that two factors keep terrestrial >>>arthropods smaller than Moose, Elephants and saurians. [quoted text clipped - 59 lines] >from any that every actually existed. Looking at Eryops, who could >have foreseen a monkey or an eagle? There is far too much speculation here and far too little comparative physiology. The physiology of scaling has been very well studied. There are numerous factors that tend to limit animal size balanced by numerous factors that tend to favor large size. The result is a balance which depends not only on internal "optimization" problems but also environmental factors like temperature and water availability, the availability and ease of access to food, and the nature of the competition.
I use "optimization" in quotes because there is far too many people look for "optimal" solutions in engineering models of animal physiology without considering the necessary costs of producing that "optimal" system. All organisms are compromises, not optima. And you need not be "best" at anything. All you need is to be better than the competition.
Clearly terrestrial arthropods could be far larger than they are today. They were in the past. Clearly terrestrial vertebrates could be larger than they are today. They were in the past. John Harshman refers to an eight foot millipede. The response was to wish for something more like a cow. When was the last time you saw a cow eight feet long? Times change, conditions change. "On being the right size" depends on the times. Incidentally, that phrase is the title of a delightful essay on scaling by Haldane and should be the starting point on the reading list for anybody interested in this subject.
Donald.Kenney@gmail.com - 07 Dec 2005 19:45 GMT ....
>1. Arthropods have exoskelatons rather than endoskelatons. That's, >I'm told, not an optimal way to build large structural elements like [quoted text clipped - 4 lines] >>build light strong structures that differ from past conventional >>wisdom.
>I strongly doubt this explanation. Some insects already have bits of >exoskeleton that extend inwards, for things such as muscle attachment. >Making more of them should not be a problem for evolution. ...
I think the problem might be a little more subtle than that. The chunk of bone that the dog has been chewing on for weeks is about 3cm in diameter with a central hole about 1.5cm. That makes the bone walls 7.5mm. A little quick spreadsheet work says that if the vertebrate bone is surrounded with muscles, nerves, tendons, et. al 1.5cm thick, the resulting appendage is 6cm in diameter and that the cross section area of the muscle etc is a smidge over 21 square cm.
If we have an exoskelaton, it probably takes the same amount of "soft" tissue to make the thing work. That means a rod of muscle, tendon, blood vessels, etc about 5.2 cm in diameter. When we cover it with the same amount of material that was in the cenral bone of the vertebrate, we find that the exoskelaton will be about 4mm thick versus 7.5mm thickness for the bone walls.
Will it be as strong as the bone? Very likely not. If we beef up the exoskelaton, it will add weight which will probably require that there be more soft tissue to support and move the heavier skelaton. This may not be all that great a trade-off when compared to simply putting the skelaton inside the critter.
r norman - 07 Dec 2005 19:58 GMT >.... >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 32 lines] >not be all that great a trade-off when compared to simply putting the >skelaton inside the critter. That explains why jumbo jets have a solid structural core with the passenger space distributed around the outside. Ocean liners, too.
deowll - 10 Dec 2005 05:17 GMT >>.... >>>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 35 lines] > That explains why jumbo jets have a solid structural core with the > passenger space distributed around the outside. Ocean liners, too. Ouch!
Richard Forrest - 10 Dec 2005 08:36 GMT > >.... > >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 35 lines] > That explains why jumbo jets have a solid structural core with the > passenger space distributed around the outside. Ocean liners, too. Yes, but the fuselage of a jet liner is not articulated via a complex joint to the fuselage of another jet liner.
I haven't done the mathematics on this, but I suspect that a limiting factor is the volume of the muscles which are needed to move articulating joints. Power is proportional to the cross-sectional area of the muscle, so muscle volume needs to increase by the cube of the power (given a muscle of fixed relative length - this is a rather hypothetical excercise).
As weight increases by the cube of the linear increase in sixe - i.e. if we double the length of an animal, the weight increases by a factor of 8, the volume of the muscles needed to move that weight increases by the fourth power - i.e by a factor of 16.
It could simply be that an arthropod runs out of space for muscles *inside* its legs segments as size increases, bearing in mind that some of that space is needed for other metabolic functions. Vertebrates, having an internal skelton, are not subject to such a limitation. If I look at the musculature of my legs, for example, it is clear that the volume of muscle is significantly greater than the volume of bone.
RF
John Wilkins - 10 Dec 2005 10:06 GMT >>>.... >>> [quoted text clipped - 40 lines] > Yes, but the fuselage of a jet liner is not articulated via a complex > joint to the fuselage of another jet liner. I'm missing something. Jet airliners *do* have a solid structural central core - at the floor of the passenger cabin. That Hawaiian airliner that lost its outer cabin wall was saved by that very structure. Had the outer skin been torn from the baggage half, the central flooring wouldn't have been able to support the weight, sure, but there certainly is internal skeletal support on most airliners.
> I haven't done the mathematics on this, but I suspect that a limiting > factor is the volume of the muscles which are needed to move [quoted text clipped - 16 lines] > > RF
 Signature John S. Wilkins, Postdoctoral Research Fellow, Biohumanities Project University of Queensland - Blog: evolvethought.blogspot.com Nihil tam absurdum quod non quidam Philosophi dixerit - adapted from Cicero
r norman - 10 Dec 2005 15:55 GMT >>>>.... >>>>>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 44 lines] >support the weight, sure, but there certainly is internal skeletal support on >most airliners. Large crustacea (lobsters) also have a similar internalized skeletal core, the endophragmal skeleton forms a relatively solid framework at the base (ventral side) of the thorax. This forms a very strong and rigid structure under which the legs attach and over which are the thoracic organs. This is very similar to the structural core at the floor of the passenger cabin. In fact I was thinking of the lobster thorax when I came across the jet liner analog. The outer shell is of both is protective, but not structural. The thorax of the lobster is also not articulated.
Yes, there are serious problems simply scaling up an exoskeleton to make a very large animal. But the suggestions given so far are not the correct way of analyzing the problem. Exoskeletons can be easily structured with appropriate ridges and fibers to resist bending and tension and hollow tubules strongly resist compression provided you eliminate buckling. That takes care of the stress problem, the first suggestion. The thickness analysis is not really correct . The strength of muscle and skeleton, the forces they can exert and support, depend essentially on their cross sectional area which scales as the square of body size. The load that must be handled depends on total body volume which scales as the cube of body size. Hence, as the animal gets larger, an increasing amount of the body must be devoted to structural support -- mouse bones are very slender while the homologous elephant bones are very stout. The percentage of a mouse body devoted to skeletal support is much smaller than the corresponding percentage of an elephant. Exactly the same considerations apply to an exoskeleton. There is no difference in the relative mass (or volume or thickness) of the skeletal vs. muscular structures in the two cases.
Probably the most important limitation for the exoskeleton is the growth pattern -- it is molted and regenerated. This, I believe, was mentioned in a different branch of this thread. Mollusc shells and vertebrate skeletons simply add new material to the preexisting structure. Evolution could solve this problem: make the exoskeleton split along specific lines (which it already does to allow the molt to occur) and then retain the old exoskeleton just growing new material at the suture lines (the ability to grow a new exoskeleton is present everywhere, now). It doesn't take that much of a change.
Articulation is another problem, but the articulations need only occur on the appendages which are far smaller than the main trunk. And, with a larger number of supporting legs, each need support only a fraction of the total load.
I think I already mentioned the problem of an open circulatory system which is severely limiting. Molluscs also have an open circulation, but the cephalopods (squid and octopus) changed to a closed circulation. Arthropods could have done similarly. Insects, instead, went the route of the tracheal respiratory system so that the circulation doesn't supply oxygen to the tissues. Even this can, in fact, be scaled up in size.
Large size arthropods would not be terribly agile, but nothing with an armor plate is. Think of Ankylosaurus or Glyptodon. Think of the giant tortoise.
The arthropods in general have been so successful at what they already do, both in water and on land, that the reorganization of the functional and developmental pattern required for large size had no particular short term advantage (and questionable long-term advantage, given the disappearance of the largest terrestrial animals). The vertebrates may not have had much of a choice of size. They were so badly outcompeted (is that a word?) in the small world that they were forced into large size. My impression is that this is a case of competitive exclusion and resource partitioning -- one group taking the small size niches, another taking the large ones.
Stanley Friesen - 11 Dec 2005 04:14 GMT > [snip] >Large size arthropods would not be terribly agile, but nothing with an >armor plate is. Think of Ankylosaurus or Glyptodon. Think of the >giant tortoise. Even this may overlook something. In a structural component, all the material outside of the stress zones is, in some sense, redundant. Thus it can be eliminated in favor of unmodified cuticle. The result, for an arthropod, would be something like a suspension bridge or skyscraper framework.
 Signature The peace of God be with you.
Stanley Friesen
John Brock - 11 Dec 2005 16:44 GMT >>Large size arthropods would not be terribly agile, but nothing with an >>armor plate is. Think of Ankylosaurus or Glyptodon. Think of the >>giant tortoise.
>Even this may overlook something. In a structural component, all the >material outside of the stress zones is, in some sense, redundant. Thus >it can be eliminated in favor of unmodified cuticle. The result, for an >arthropod, would be something like a suspension bridge or skyscraper >framework. Or the structural support might even gradually migrate to the interior, and end up looking like an endoskeleton. (As I pointed out earlier, the vertebrate skeleton started out as an exoskeleton). But even if that happened I don't think the resultant animal would end up looking much like a vertebrate, and that what makes this exercise interesting to me. What would a really big arthropod that never had to face competition from tetrapods look like?
 Signature John Brock jbrock@panix.com
r norman - 12 Dec 2005 02:49 GMT >>>Large size arthropods would not be terribly agile, but nothing with an >>>armor plate is. Think of Ankylosaurus or Glyptodon. Think of the [quoted text clipped - 13 lines] >exercise interesting to me. What would a really big arthropod that >never had to face competition from tetrapods look like? The endoskeleton of the vertebrates does not derive from the outer protective casing migrating inward. It is a completely separate system.
John Brock - 12 Dec 2005 16:15 GMT >>>>Large size arthropods would not be terribly agile, but nothing with an >>>>armor plate is. Think of Ankylosaurus or Glyptodon. Think of the >>>>giant tortoise.
>>>Even this may overlook something. In a structural component, all the >>>material outside of the stress zones is, in some sense, redundant. Thus >>>it can be eliminated in favor of unmodified cuticle. The result, for an >>>arthropod, would be something like a suspension bridge or skyscraper >>>framework.
>>Or the structural support might even gradually migrate to the >>interior, and end up looking like an endoskeleton. (As I pointed [quoted text clipped - 3 lines] >>exercise interesting to me. What would a really big arthropod that >>never had to face competition from tetrapods look like?
>The endoskeleton of the vertebrates does not derive from the outer >protective casing migrating inward. It is a completely separate >system. True, but that doesn't really change the point. The structural material -- bone -- in vertebrates was originally used externally, as a carapace, but ended up being used internally, as an endoskeleton. It's possible something similar could have happened with our hypothetical large land arthropods. With land animals I kind of think there *would* have needed to be a "migration", with internal projections originally used as muscle attachment points gradually taking over structural function. All speculation of course, but I don't see any reason it couldn't have happened that way.
 Signature John Brock jbrock@panix.com
r norman - 12 Dec 2005 16:30 GMT >>>>>Large size arthropods would not be terribly agile, but nothing with an >>>>>armor plate is. Think of Ankylosaurus or Glyptodon. Think of the [quoted text clipped - 27 lines] >taking over structural function. All speculation of course, but >I don't see any reason it couldn't have happened that way. All of these discussions about "might have been" are interesting but so speculative that I am not sure how far to push things. Certainly insects "could have" modified their exoskeleton to include a far more internalized system like the endophragmal skeleton of the crustacea. And insects "could have" used the cellular biochemistry that produces hard and rigid chitin to make an internal skeleton. And insects "could have" developed a closed circulatory system and beefed up the tracheal respiratory system with more effective ventilation. I still think that large arthropods are conceptually possible, at least for the purpose of writing exotic "science" fiction and fantasy stories and movie scripts. Biologically it is extremely improbable.
It certainly is clear that vertebrates had a distinct advantage in the size race with their internal bony skeletons that evolved for whatever reason in the water but gave them an enormously valuable pre-adaptation for structural support on land. In that competitive environment, terrestrial vertebrates took the large-size route while insects and arachnids and other arthropods took the small-size route. Both lead to success and it is hard to tell which group is the ultimate "winner" (whatever that might mean).
John Brock - 13 Dec 2005 15:30 GMT >All of these discussions about "might have been" are interesting but >so speculative that I am not sure how far to push things. Certainly [quoted text clipped - 16 lines] >Both lead to success and it is hard to tell which group is the >ultimate "winner" (whatever that might mean). The thing is, it's not just a matter of "could have," there is a "would have" element as well. Without competition from vertebrates -- which I agree started out just better adapted for the large land animal niches -- arthropods would definitely have done *something*.
One possibility is that they would have ended up as nothing more impressive than moderately larger and more sophisticated versions of the same sorts of arthropods we've already come to know and love. That seems to be what you are arguing for when you suggest that anything else is "extremely improbable," and for all I know you may be right. But personally I think it is rather likely that they would have evolved into new forms which would have been profoundly different than any that have ever actually appeared, and I strongly suspect -- since the niches do exist -- that some of them would have ended up much larger and more active than the real-life sprawling 5 foot sea scorpion that started this thread.
 Signature John Brock jbrock@panix.com
deowll - 18 Dec 2005 05:20 GMT >>>>Large size arthropods would not be terribly agile, but nothing with an >>>>armor plate is. Think of Ankylosaurus or Glyptodon. Think of the [quoted text clipped - 17 lines] > protective casing migrating inward. It is a completely separate > system. You are thinking notochord. The first boney material was in the skin. In the boney headed fish it stayed there to some degree. If we were descended from them our heads would look rather different.
r norman - 10 Dec 2005 16:08 GMT >> >.... >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 57 lines] >look at the musculature of my legs, for example, it is clear that the >volume of muscle is significantly greater than the volume of bone. Not really correct. I treated scaling in another post on this thread. As I pointed out there, exactly the same problem of scaling between muscle and skeletal structures applies to internal skeletons: as the linear size increases, the proportion of body mass devoted to structural support must increase. Note: if you look at the legs of a small animal like a mouse, you find that the volume of muscle compared to bone is even larger.
Muscle force is proportional to the cross-sectional area. The distance muscle can move is proportional to the length. The work that muscle can do or the energy required to do the work is proportional to the volume. But the energy available from cellular metabolism is also proportional to the volume. Power is the rate of expenditure of energy and doesn't really apply. There is no fourth power relation involved.
Richard Forrest - 11 Dec 2005 17:31 GMT > >> >.... > >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 73 lines] > energy and doesn't really apply. There is no fourth power relation > involved. I'm refering to "power" as a mathematical term, not a mechanical term!
The force exerted by a muscle is proportional to its cross-sectional area. The mass of the muscle is proportional to its volume. So if we double the length of the whole animal, we increase the volume, and hence the weight of the whole animal by the cube of the increase, have to increase the cross-sectional area of the muscles needed to move the weight around by the cube of the increase, and therefore increase the volume - and hence the weight - of those muscles by the fourth power.
This is one of the reasons why large animals tend not to lie down, tend walk stiff-legged, and don't engage in activities such as jumping which need to lift the whole weight of the body off the ground. It's to minimise the proportion of the muscle mass needed simply to move the body weight.
RF
r norman - 12 Dec 2005 02:54 GMT >> >> >.... >> >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 89 lines] >minimise the proportion of the muscle mass needed simply to move the >body weight. I still don't understand where the fourth power comes in. You are correct in repeating what I said, that the force available increases as the square of body size while the force needed increases as the cube of body size. That simply means that muscle and bone scale muscle and bone cross sections increase as the cube of body side, hence the diameter or thickness of muscle or bone increases as the 3/2 power of body size. As a result, larger animals have a larger percentage of their body mass devoted to support than smaller ones. Moving more sluggishly (with less acceleration) means needed less muscle force and reducing the demand for larger muscles.
josephus - 12 Dec 2005 07:23 GMT >>>>>>.... >>>>>> [quoted text clipped - 102 lines] > Moving more sluggishly (with less acceleration) means needed less > muscle force and reducing the demand for larger muscles. I saw earlier a remark that giant creatures must move stiff legged and slowly. How does this knowledge match Robert Bakker's hot blooded dinos. He has medium sized monsters climbing trees, and fighting predators. I have read the book by Bakker. I found his arguments compelling. josephus
r norman - 12 Dec 2005 13:15 GMT >I saw earlier a remark that giant creatures must move stiff legged and >slowly. How does this knowledge match Robert Bakker's hot blooded >dinos. He has medium sized monsters climbing trees, and fighting >predators. I have read the book by Bakker. I found his arguments >compelling. > josephus I haven't read the book. Just how big are the tree-climbers? Modern bears climb trees. Were the "medium sized" monsters significantly bigger than that? And "stiff legged with limited agility and mobility" needn't translate into slow. The predator-prey battles between behemoths were between two equally stiff legged animals of limited agility.
deowll - 18 Dec 2005 05:24 GMT >>>>>>>.... >>>>>>> [quoted text clipped - 120 lines] > compelling. > josephus A lot of people think that he over stated his case for speed for the really big animals. That the smaller ones were very fast and did climb trees isn't in question for some of them. My guess is a t-rex was as fleet footed as it needed to be.
John Brock - 18 Dec 2005 23:41 GMT >"josephus" <dogbird@earthlink.net> wrote in message
>> I saw earlier a remark that giant creatures must move stiff legged and >> slowly. How does this knowledge match Robert Bakker's hot blooded >> dinos. He has medium sized monsters climbing trees, and fighting >> predators. I have read the book by Bakker. I found his arguments >> compelling.
>A lot of people think that he over stated his case for speed for the really >big animals. That the smaller ones were very fast and did climb trees isn't >in question for some of them. My guess is a t-rex was as fleet footed as it >needed to be. I would rephrase that slightly to say that T-Rex was as fleet footed as it *could* be. In "Predatory Dinosaurs of the World" Gregory Paul argued that "if an animal looks fast, it's fast," and then went on to explain why T-Rex "looks fast" to him. I thought his argument was persuasive, but it doesn't really answer the question "how fast?" The cheetah (and to a lesser extent other big cats) is highly optimized for speed, because its lifestyle is such that the faster it can run, the better.
But that doesn't mean it's ever going to hit 120mph, however big a selective advantage that would bring! Likewise I think that T-Rex was also optimized for speed, and that the faster it could run the more food it could catch. But that still doesn't answer the question. For all we know with an animal that size evolution may hit the wall at 15mph! (Still faster than a lot of people of course, and remember that the animals it was chasing would also be large, and thus would face similar constraints). But it doesn't do to underestimate Mother Nature, or imagine we can anticipate all of her clever devices, so my guess would be a top speed in the 20-25mph range, or 30mph at an absolute upper limit. (Note that Paul considered 30mph to be the lower limit!).
 Signature John Brock jbrock@panix.com
Richard Forrest - 12 Dec 2005 09:10 GMT > >> >> >.... > >> >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 100 lines] > Moving more sluggishly (with less acceleration) means needed less > muscle force and reducing the demand for larger muscles. If the linear dimensions of the muscle is doubled, the force it is able to exert goes up by the square - i.e. x 4. However, the weight of the muscle goes up by the cube - i.e. x 8. So if the muscle is involved in lifting its own weight, it has to increase its cross-sectional area by more than simply the cube - i.e. to a power of 3+. At small scales this factor is insignificant, but as animals become larger, the proportion of their bodyweight given over to musculature becomes an increasingly significant proportion of their total weight. Large animals use behavioural and biomechanical strategies to restrict this by either shortening the limbs so that the muscles needed to lift the body are shorter and rounder - hippos and rhinoceroses are examples of this - or by avoiding the circumstances under which there is a need to lift their own body weight, and locking the knee joint when walking - giraffes and elephants, for example. If you watch a film of elephants, you will see that although the smaller ones bounce along, the bigger ones walk stiff-legged. The knee-joint is locked before the foot hits the ground, so that the muscle force which goes into lifting the body when walking is minimised.
RF
r norman - 12 Dec 2005 13:07 GMT >> >> >> >.... >> >> >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 119 lines] >so that the muscle force which goes into lifting the body when walking >is minimised. After sending my message I realized I was wrong. I saw where the fourth power comes from. I also went through the same arguments you are now presenting about the limited agility and range of movement that the muscle and joints provide.
In any event, these same calculations apply equally to vertebrates with internal skeletons and to arthropods with external skeletons. I don't envision a large arthropod having a uniform outer shell, but rather having distinct heavy ridges and thickenings longitudinally arranged for the main structural support and a rather thin protective shell covering the rest of the circumference.
Richard Forrest - 12 Dec 2005 17:53 GMT <snipped>
> After sending my message I realized I was wrong. I saw where the > fourth power comes from. I also went through the same arguments you [quoted text clipped - 7 lines] > arranged for the main structural support and a rather thin protective > shell covering the rest of the circumference. This "fourth power factor" formed one of the main arguments in support of a new theory of dinosaur extinction hatched in a pub over several drinks during a conference a few years. It goes like this:
As animals increase in size, the amount of muscle mass increases exponentially. For mammals it peaks out at around 10 tons. So large sauropod dinosaurs, some of which have been estimated as weighing 80 tons (though I suspect that such estimates were *also* the result of late-night drinking sessions) were finding a way around this limit.
Feeding is no particular problem. The long neck provides a large area of grazing, so they can feed from a large volume of vegetation with very little effort. Walking can be slow and stately, never putting much dynamic load on the joints. The problem arises (as it so often does) with sex.
When a male sauropod is humping a female sauropod, his weight has to be supported by her hind legs, and if they are already loaded to close to their limit. So either the male is much smaller than the female or he has a long, steerable penis which can be inserted from a side-to-side stance. (Having a "bit on the side", as we say in England). However, such speculations are unsupported by any evidence, which leaves us with a third option: large sauropods were all female, and reproduced parthogenically. This idea is supported by some evidence - turkeys occasionally do this, and they are, after all, related.
This is fine, so long as there are no major changes in environment.
We now have to take thermal control into account.
Large sauropods were passive homotherms, and because of their enormous size could maintain a very stable body temperature, responding only very slowly to changes in the temperature of the environment. However, although such changes were slow, they were hard for the animals to control. They could not increase metabolic rates during cold weather by increased activity (remember the "fourth power" factor reducing their gait to a slow and stately progression), so if the season turned cold, they would need to migrate to warmer climates to compensate.
Similarly, during hot seasons they would find it very hard to lose heat, and would need to head for cooler climes during the warmer part of the year. So they became locked into a seasonal migration between tropical and temperate latitudes simply in order to maintain a constant body temperature. Large herds of giant sauropods moving across a landscape and eating every scrap of vegetation has a profound effect on the ecology.
This is where we have to consider the ecology of large sauropods.
Sauropods had, as we now know, bony plates and osteoderms on their backs, and a rough, rugged skin. This would have provided the nooks and crevices needed for small plants to grow, these would in turn attract insects and other invertebrate feeders, and thes would attract the small mammals and pterosaurs feeding on them. So the sauropod was the centre of a whole ecological system in much the same way as an oak tree is today.
Now plate tectonics comes into play.
During much of the Mesozoic, the continents were arranged so that the migration routes from the tropics to the north and south was relatively easy, and involved no water crossings. Towards the end of the Cretaceous however, the continents fragmented, and these routes became barred by stretches of deep water. So the sauropods, unable to evolve into new forms to overcome this problem because they were both parthogenic and long-lived, died out, and their demise brought a whole slew of ecological systems crashing down with them, bringing radical changes to climate and rainfall patterns, and triggering a mass extinction event.
Curiously, this speculation was witnessed by a BBC researcher working on an proposal for "Walking with Dinosaurs". A small detail - the pterosaurs nesting on the backs of the sauropods - actually made it into the final version.
RF
r norman - 12 Dec 2005 18:21 GMT ><snipped> > [quoted text clipped - 84 lines] >pterosaurs nesting on the backs of the sauropods - actually made it >into the final version. My guess is that quite a few more than "several drinks" were associated with that discussion. I would be interested in more details the problems of scaling applied to penile erection, not to mention the guidance problem associated with the notion of "steerability".
There is at least one problem with your analysis: that of large sauropods moving across the environment eating every scrap of vegetation. The fact is that those large sauropods, although they do eat quite a lot of stuff, have a far lower total metabolism than the same biomass distributed among much smaller animals. That is, the same biomass in sheep-sized grazers would eat far more; in rabbits and mice much more still, and in insects they would quickly devastate the landscape the way migratory locusts do.
Also there are two very effective ways to increase metabolic heat production during cold weather that does not involve rapid locomotion. Every mammal knows about shivering and non-shivering thermogenesis. The latter is a biochemical pathway that burns glucose but uncouples electron transport from ATP production so the process is energetically absolutely inefficient: the only product is "waste" heat. Of course, that is exactly the desired effect. Even some insects know how to shiver in a way, buzzing their bodies to produce enough metabolic heat to increase their overall metabolic rate so they can fly in cold weather.
Richard Forrest - 12 Dec 2005 19:58 GMT > ><snipped> > > [quoted text clipped - 90 lines] > mention the guidance problem associated with the notion of > "steerability". If you look at the way the tail of a large dinosaur attaches to the body, and the massive musculature anchored to the base of the tail (including some of the locomotor musculature), sex becomes a bit of a problem for a mammalian type of "humping". I believe that Alan Charig gave a talk about dinosaur sex which explained how this problem was circumvented, complete with demonstration (of posture at least) with a female colleague, but this was before my time. I don't know if this appeared in the literature.
> There is at least one problem with your analysis: that of large > sauropods moving across the environment eating every scrap of [quoted text clipped - 4 lines] > and mice much more still, and in insects they would quickly devastate > the landscape the way migratory locusts do. True, and according to Kent Steven's analysis of neck mobility, the larger brachiosaurs would not have been able to feed easily from something at ground level. Dipolodocids apparently could, but they didn't reach the same enormous size.
On the other hand, the pounding of hundreds of meter-diameter feet across a landscape wouldn't do the vegetation much good.
> Also there are two very effective ways to increase metabolic heat > production during cold weather that does not involve rapid locomotion. [quoted text clipped - 6 lines] > to increase their overall metabolic rate so they can fly in cold > weather. Somone raised this, and someone else (the identities escape me for now) stated authoritatively that reptiles don't shiver. I don't know if this is universally true, but friends of mine who keep reptiles confirm this. If they get cold, they simply get more and more torpid. I believe that some boas "shiver" to generate heat when incubating their eggs, but apparently don't if they get cold at other times. However, considering the volume to surface area ratio of an 80 ton sauropod, I'd guess that getting rid of excess heat would be more of a problem than chilling.
RF
deowll - 18 Dec 2005 05:36 GMT >> ><snipped> >> > [quoted text clipped - 139 lines] > > RF Their guts contained huge fermentation chambers that would have generated heat. Unless things got rather cold I doubt if they had much of a problem. Getting enough food would have been the problem.
deowll - 18 Dec 2005 05:33 GMT > <snipped> > [quoted text clipped - 37 lines] > > This is fine, so long as there are no major changes in environment. They may have mated when small and retained sperm for later egg fertilization or they many not have been as near their weight limit as some people think.
> We now have to take thermal control into account. > [quoted text clipped - 44 lines] > > RF deowll - 18 Dec 2005 05:10 GMT >> >.... >> >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 59 lines] > > RF Some arthropods us hydrolics in their legs.
NashtOn - 18 Dec 2005 12:29 GMT >>>>.... >>>> [quoted text clipped - 63 lines] > > Some arthropods us hydrolics in their legs. To the OP:
All you would need to do is increase the distance between the axis of movement and the insertion of the muscle to increase generation of rotary torque. Cross section and length of muscle notwithstanding, a minute increase in leverage has a profound effect.
Furthermore, any increase in PCSA has hardly any effect on intrinsic muscle properties, even though it has dramatic effects on extrinsic muscle properties.
http://muscle.ucsd.edu/musintro/pcsa.shtml
My point is that production of force is more a function of the ability of the muscle to produce rotary torque (the further the point of insertion, the better) than PCSA.
http://muscle.ucsd.edu/musintro/ma.shtml
To complicate matters, muscle fibers come in very different varieties as some muscle tissue is made for speed, others for endurance and all the intermediaries.
To further complicate matters, optimum recruitment of motor units would have to be considered. IOW, you can have a critter with very little muscle but with a very efficient CNS/PNS that "evolved" into being able to optimize motor unit recruitment.
To further complicate matters, you would have to know exactly what type of niche said creature occupied in the temporal context you were discussing. Was it at the top or bottom o the food chain? Was it arboreal or did it live where trees were sparse? If it were at the bottom of the food chain and needed to run fast to avoid predators, it would need fast glycolytic fibers, it would have to have minimum mass and would need to sleep many hours to replenish its energy.
Now, draw your own conclusions concerning the advantages of an endoskeleton vs an exoskeleton in your simple discourse, when most of the data to prove your points is lacking.
What is amazing is how you evolutionists can simplify something as complicated as muscle function and draw conclusions based on PCSA for example, without considering other important factors and then add some complicated words to the mix to sound intelligent, when in fact, you have absolutely no idea whatsoever of what you're talking about.
Nicolas
Stanley Friesen - 07 Dec 2005 21:15 GMT >I think the problem might be a little more subtle than that. The chunk >of bone that the dog has been chewing on for weeks is about 3cm in [quoted text clipped - 12 lines] > >Will it be as strong as the bone? Very likely not. Actually, much depends on the structure. In all likelihood the stress zones would be thicker, and have ridges oriented parallel to the main stresses, and the remaining parts would be thinner, or even reduced to just cuticle. The result might well be something effectively as strong as bone at a similar weight.
 Signature The peace of God be with you.
Stanley Friesen
deowll - 10 Dec 2005 05:16 GMT > .... >>1. Arthropods have exoskelatons rather than endoskelatons. That's, [quoted text clipped - 32 lines] > not be all that great a trade-off when compared to simply putting the > skelaton inside the critter. Which might make something the size of a Bronto out of the question.
I still think with a few adjustments a future bioengeers might get something as large as the largest turtle to work at least as well.
In other words I do agree that bigger animals do better with an endoskeleton and this one reason that larger arthropod species are uncommon but I consider molting to be an even larger problem and has limited their ability to compete in the 20 to maybe 200 pound range.
rms - 01 Dec 2005 20:56 GMT > A unique, complex trackway has been discovered in Scotland: it was > made roughly 330 million years ago by a huge, six-legged water > scorpion that was about 1.6 m long and a metre wide. To my knowledge, > this is not only the largest terrestrial trackway of a walking > arthropod to be found so far, but is also the first record of > locomotion on land for a species of Hibbertopterus (Eurypterida). _Australia: The Four Billion Year Journey of a Continent_, has a nice picture of Eurypterid tracks which the caption says are 400Myr, "tracks up to 20cm wide in Murchison Gorge suggest that the maximum length of these animals was a little less than a metre, though younger fossils twice that size have been found elsewhere."
rms
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