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Re: Bipedalism



At 05:13 PM 02/04/98 -0600, Rob Meyerson wrote:

>I went with the idea that bipedalism has it's advantages for cursorial critters and can be seen as a cursorial adaptation; I then went on to suggest that theropod bipedalism evolved on the ground, before they moved into the trees.

I think this is only partly true, and the reason for this is the nature of the spinal column.  This is where we run into one of the big differences between mammals and dinosaurs/birds.  If you have a spinal column capable of a considerable degree of dorsoventral flexion and extension, bipedality is certainly NOT going to be an advantage for cursoriality.  Instead, you would be best as a quadruped like a cheetah, which can arch its spine to bring its hind legs well forward during running and thereby increase stride length far beyond what would be possible for a similar-sized running biped.  This may be why there are so few bipedal terrestrial mammals, and those that do exist - with ourselves as the only exception I can think of - hop rather than run (kangaroos, jerboas, the springhaas).  And, of course, we are almost certainly bipedal as a result of having evolved from arboreal brachiators, a most unusual development (perhaps the extinct ground sloths, and some of the subfossil Malagasy lemurs, shared the same pathway). [This, BTW, is one reason I find it hard to swallow the sci-fi premise that the humanoid body shape is the likeliest one for intelligent aliens!]

In birds, however, we have an almost inflexible spinal column, and for them bipedality has certainly turned out some pretty fast runners - an ostrich is one of the fastest land animals going.  But whether bipedality evolved as a cursorial adaptation is another matter - it could have evolved because it was advantageous to have the hands free to perform other roles, or for rearing to reach higher vegetation.  About all I would care to say is that bipedality was not evolutionarily possible until the spinal column lost much of the LATERAL flexibility seen in early tetrapods, so that it related to a shift in posture and gait - but these would have to have evolved first, making bipedality a structural option whose initial advantage may have had nothing to do with locomotion (or, to give a nod to George, with terrestrial locomotion).

>To take a nod to BCF, I then suggested the following model: Bipedalism evolves on the ground; >these "prototheropods" discover an advantage to life in the trees, and begin to adapt to that >lifestyle; a part of that adaptation is longer arms, to help grasp the branches; these longer arms >are now preadapted to become wings.  These guys, then, are the ones that evolve into true >theropods.

I doubt that there is any way to establish whether bipedality evolved first on the ground or as a climbing adaptation, for the reason suggested above - namely, that it may have evolved for reasons unconnected with locomotion, reasons that could apply either on the ground or in trees (eg prey handling).  One of my big problems with BCF (which has its attractions as a concept) is that it assumes that you can make this sort of distinction.  I haven't the faintest idea how you would tell from structure alone, assuming you had the fossil, if the first bipedal archosaur lived in trees, low bushes, rock piles, open country or all of the above, and from a purely phylogenetic standpoint of course it doesn't matter.

>I acknowledge that bipeds can adequately function in trees, with some modification of their form.  >Having claws or wings certianly helps things along.  I merely suggest that the bipedalism *trait* is >most likely to develop on the ground.

I would point out that any biped that does NOT have claws or wings is going to be a highly derived form indeed!  In fact I cannot think of a single bipedal tetrapod (leaving aside certain burrowing lizards and pythons, which hardly qualify as bipeds) that has entirely lost all pectoral skeletal elements while retaining the pelvic ones.

In fact, unless you believe that bipedality evolved at (as it were) a single bound, the first archosaurs to assume bipedal postures and gaits must have had well-developed, fully-functional forelimbs.  These could well have been used for climbing quite early in the evolution of bipedality - in fact you could argue (though again I know no way of proving this) that the more advanced a biped becomes in terms of reduction or modification of the forelimb elements, the less well-adapted it might be for hauling itself up into a tree.  I can see bipedality evolving either on the ground or in trees, with early generalized bipeds having considerable potential to shift either into or out of trees and back again.  Thus I don't see bipedality as an argument either for or against an arboreal habit in its first possessors.

>>Then why do monkeys and apes retain a normal walking ability?  Well I
>>suppose hopping/leaping will only become obligatory if the animal returns to
>>the ground only very rarely, and then it may not happen (only some of
>>lorris's, potto's etc leap/hop, and many apes are too big to risk it).
>
>Firstly, I would point out that most of the above are "temporarily bipedal."  When chimps or gorillas travel in a casual walk, they are quadrepeds, using all four limbs for motion.  These critters stand only occasionally, and only for brief periods of time.  Basically, their skeleton is not designed for constant bipedalism.

But gibbons ARE obligate bipeds on the ground - I have never seen one walking quadrupedally.  Bear in mind that one of the biggest anatomical shifts in primate locomotion was not coming out of the trees (which quadrupedal primates like baboons, patas monkeys and ring-tailed lemurs have managed quite nicely) but shifting from running/leaping to brachiation.  Brachiation has produced major modifications in the structure and proportion of the limb elements, and I would speculate that in primates at least (not in sloths, for example) it involved loss of quadrupedality at an early stage.  It strikes me that the larger apes are secondarily quadrupedal on the ground as a function of increased size; the fact that they walk on their knuckles, a most unusual type of gait, suggests to me that their ancestors lost the ability to move quadrupedally on the ground and that this had to be re-evolved.

--
Ronald I. Orenstein                           Phone: (905) 820-7886
International Wildlife Coalition              Fax/Modem: (905) 569-0116
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