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echo: evolution
to: All
from: William Morse
date: 2004-05-14 06:02:00
subject: Re: Species selection, Wa

john_SPAM{at}wilkins.id.au (John Wilkins) wrote in
news:c7o7jd$2jr0$1{at}darwin.ediacara.org: 

> William Morse  wrote:
> 
>> john_SPAM{at}wilkins.id.au (John Wilkins) wrote in
>> news:c7hmdm$i0l$1{at}darwin.ediacara.org: 

> As Eldredge admits, in his _Macroevolution_, the reproduction is not,
> really, reproduction. That is why the term "moremaking" is introduced
> - cars are made more of, Xeroxes are made more of, but what is
> lacking, what is necessary for selection, is the idea of hereditable
> characters which correlate with economic success. Do species reproduce
> in ways that are successful relevant to their properties?

Thnaks for the clarification, At first I thought Eldredge might have been 
objecting to the idea that the number of species has increased over time, 
which has been a bone of contention. As I understand it now, his 
objection is your objection - that even though species create other 
species, those other species do not inherit anything that affects their 
success as a species. 
 
> As to the sampling of gene frequencies - for a start, at least some
> models of speciation (Mayr's peripatric, White's stasipatric, and, I
> would argue, sympatric) do *not* sample identical alleles and
> freqencies. Imagine, as a thought experiment, a species that is
> identically divided, and the evolution of which is parallel in each
> half. Are they (causally speaking) different species? Obviously not if
> they will freely breed and not change alleles when in sympatry again.
> 
> Of course it is highly unlikely that isolates would evolve in parallel
> even if they were identical to begin with, but leave that to one side.
> 
> Consider this illustration. I sample a bucket of pingpong balls of
> different colours. Is the sample set the same as the ratio of colours
> in the original bucket? Obviously that is dependent on the size of the
> sample. Suppose it isn't - has the sample "inherited" its colour
> frequencies? Suppose it is - can we say it *did* inherit the
> frequencies, or is it a matter of chance? I say chance, and this
> applies equally to allopatric speciation.

I understand the concept of sampling. The problem is that sampling is a 
population genetics concept, and that may be  irrelevant to the question, 
which is an ecological niche concept. Remember (I know that you know the 
history a minimum of 16.3 times better than I do) that Darwin and Wallace 
came up with the concept of evolution without any knowledge of genetics. 
The fact that they did this is an argument that the genetics is not 
necessary to the argument. It adds to our understanding in many cases but 
in other cases (such as this) is only obfuscating. My point was that, at 
the time of their geographic isolation, both populations shared the same 
niche. Until the niche changes, whether or not they share the same 
genetics (we know that thanks to drift they will not), both populations 
will remain substantially identical (with respect to the niche) other 
than that they may no longer be able to interbreed (I can explain how 
this can occur based on gene duplication, but I would prefer not to waste 
the time at this point). I understand quite well that sympatric 
speciation requires niche differentiation, which is why I noted the 
difference between sympatric and allopatric speciation, which you appear 
to not have understood based on the following paragraph: 

 
> I think there is nothing about allopatric speciation that suggests
> that the frequencies *are* inherited, and all other speciation modes
> rely upon them *not* being. Hence the causal relationship between
> parent and progeny is insufficiently fixed for selection. And hence it
> is a sorting process. We are being misled by language here.


As I explained above, the frequencies are a red herring. What is 
important is the niche, and that is initially conserved in allopatric 
speciation but not I think in the other cases. Sympatric speciation would 
seem to require niche differences. 

 
> That there is comeptition between organisms, or even between
> populations, is not objectionable to me. That there is anything about
> *species* that allows them to be in competition (again, I wearily
> state, *as species*) is. So far I can only see species competing when
> they are a single population, or at best a cohesive metapopulation. 
 
Others have answered this, and I think in general we have pretty much 
flogged the subject to death. I understand that you do not think the fact 
that a species consists of individuals that fly means that I can include 
the species in a set of "flying species". So even though I can 
demonstrate that "flying species" are more likely to colonize islands and 
undergo subsequent speciation, this will not imply species selection. I 
disagree on the grounds of utility, but your position appears to be 
logically consistent, and you show no signs of being convinced by any of 
the rest of us who have argued against that position. 

Note that Wilson has argued that species that have mechanisms that damp 
out population fluctuations (which is at least selection at the 
population level, since individuals cannot exhibit fluctuations in 
population size) will tend to go extinct less frequently than species 
with large swings in population size. It is not clear to me that this 
characteristic will be reliably inherited by daughter species, so I am 
reluctant to press this issue too hard.



Yours,

Bill Morse
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