Subspecies

[Cris Guppy & Aud Fischer]

James,

Perhaps I over reacted, but I don't think so given the way you presented
your views. I do tend to get a bit intense at times however, and I apologize
if my response seems overly agressive.

You are concerned with the recognition of subspecies that are not
consistently phenotypically distinct. That does occur on a regular basis,
but is dealt with through normal taxonomic revision processes. Every so
often the available subspecies and species names for a group are reviewed,
and (in theory) the revisor documents which ones are consistently
phenotypically and presumably genotypically distinct, and which ones are
not. This is a constant and normal process: A set of apparently well defined
taxa are named, further research occurs that shows that some are not well
defined after all, and some taxa are synonymized with others. I see no
problem with this, although reasonable care taken in the initial description
of a taxon to ensure that it really is a well defined unit separate from
related ones. And as Norbert mentioned, all too often the revisor simply
synonymizes without documentation or rationale. You do this yourself with
your blanket condemnation of the "Systematics of Western North American
Butterflies", by which you come close to implying that all the new
subspecies should be rejected. I am quite sure that this is not what you
meant, but that is what your words say. I strongly suspect that you (given
where you live) do not yourself have sufficient knowledge of most of the
taxa involved to provide a well reasoned rejection of the new taxa, and
criticize the book on the basis of second hand opinions. I certainly do not
agree with everything in the "Systematics" book, but where I disagree I will
publish a careful rationale for my alternate view and not broadcast a public
blanket condemnation.

You provide Euphydryas as an example of where a number of recognized
subspecies are not consistently different from other recognized subspecies,
with the criticism based on the opinions of friends who are presumably well
versed in the taxa involved. This is certainly the case in Euphydryas, but
the solution is not to make vague undocumented assertions and condemn the
use of the subspecies concept (yes I know, a broadscale condemnation is not
what you really meant, but it IS what you said). The solution is to publish
a reasoned rearrangement of the taxa involved, with documented synonymies.

Some butterflies, such as Parnassius smintheus (yes, I accept the split from
P. phoebus because the split was made using a wide range of well documented
evidence), have almost every population phenotypically distinct (on average)
from every other population. I believe that this is the type of variation
that you cite as resulting in "oversplitting", and that can indeed happen.
This can be taken care of through a careful analysis of the variation that
exists, and the selection of appropriate characters to separate groups of
populations that lack significant gene flow between them (ie, your concept
of a subspecies). The problem with oversplitting is resolved through a
carefully rationalized revision, with appropriate synonymies, not with a
blanket condemnation of the subspecies concept.

Just to continue to keep you on your toes, I did NOT agree "that some people
do throw names out there in the case of vanity that are unjustified".  I
simply agreed that sometimes vanity is the motivation for naming a taxon,
but as far as I am aware most people (vain or not) try to ensure that they
are naming a valid taxon because otherwise it will be synonymized and sink
into obscurity. I am not concerned with motivations for naming (beyond
normal human curiosity), I am only concerned with whether the taxon is a
discrete entity or whether it is simply an inadvertent redescription of a
pre-existing taxon.

I am not aware of Ken's "alternate approach", but from your description it
appears that instead of a Latin subspecies name an English (or presumably
other language) place name would be used. Within the existing system this
could be accomplished by simply naming all subspecies after their type
locality or distribution, which is frequently done. Ken's approach could
work, but since the subspecies approach is "not broke" why try to fix it?
[Ken will hopefully correct any of my misconceptions of his approach]. My
personal opinion is that generally a taxon should be named after one of its
characteristics (type locality, distribution, foodplant, habitat,
morphological character, behaviour, etc), however I also have no objection
to naming a taxon after a person. And I do not object to others using
different criteria for selecting names. The taxon name is just a lable, and
that is how it has been treated through most of post-Linnaean taxonomy
[think of all the species that have been named after figures or places in
Roman or Greek mythology with no obvious relationship to the biological
entity]. But there is no reason that everyone has to have the same approach
to naming taxa, as long as the basic ICZN rules are followed to prevent
chaos.

Anyway, I think this is probably all I will say on this subject through
"leps-l". I think I have provided enough of my perspective, and there is no
point droning on. I am quite happy to continue the discussion privately
should you wish.

Cris.

-----Original Message-----
From: DR. JAMES ADAMS <JADAMS at em.daltonstate.edu>
To: leps-l at lists.yale.edu <leps-l at lists.yale.edu>
Date: January 27, 2000 11:36 AM
Subject: Re: Subspecies


>Chris,
>
>> I have never understood why those who dislike the concept would care
>> enough to attack those that do find subspecies meaningful.
>> Unfortunately I still don't.
>
>You are way too sensitive!  I'm not attacking those who find
>subspecies meaningful; indeed, I'm not even necessarily saying
>that subspecies have no place in taxonomy at all.  But there *are*
>those who have, as far as I can tell, named "subspecies" with little
>or no reason other than the entity looks a *little* different, or its
>simply found in a different place.  *This* is what I object to -- no
>*biologically relevant* reason for the name.  Contrary to what may
>have come across from my previous tirade, I use subspecific
>names myself, for entities which seem to be good vicariant entities.
>More about this below.
>
>QUESTIONS: (1) What do you mean by "truly something
>> different"? Obviously subspecies will not have major differences by
>> definition, they are all the same species by definition.
>
>Agreed, to a point.  But this is my main objection in the first place.
>Unless I am mistaken, subspecific names *should be* applied to
>represent an entity which are biologically distinct in some way, and
>may potentially have it's own evolutionary path to follow.
>Otherwise, the names are nothing other than a human
>convenience, and may not represent a truly *distinct* taxon.
>
>(2) What do
>> you mean by "truly genetically differentiated"? How different is
>> "truly" genetically differentiated?
>
>There *is* no standard, obviously!!  Otherwise, we wouldn't be
>having this discussion!!
>
> (3) Why should subspecies mean
>> "anything important evolutionarily"? Do species mean anything
>> important evolutionarily?
>
>These are the only taxa which potentially *do* mean something
>important evolutionarily.  I'm sorry if I was using generalized jargon,
> but I thought it was pretty clear that I was talking about "natural"
> ("real"?) entities here.  Species within genera, families, etc. care
> nothing about what's happening evolutionary to other specific
>members of those higher level taxa -- each species has its own
>evolutionary potential, independent of others.  Genera and all higher
>taxa are purely a matter of convenience for workers to indicate
>some level of relatedness, and clearly subjective.  Species and
>subsets thereof are the only entities upon which evolution can
>actually work, because these are the entities which are, by
>definition, still genetically unified in some way.  This is what I was
>referring to when I said "important" -- if the subspecies concept is
>going to be useful, it should apply to subsets of species which,
>although not distinct enough to be called species, have some
>genetic potential independent of other subspecies.  I'm sorry if it
>sounds to general, but I'll leave it at that for now.
>
>(4) British Columbia has been almost 100% recolonized by
>> butterflies after the last glaciation, because it was nearly
>> completely covered by ice. The geographic arrangement of subspecies
>> provides information regarding the location of populations outside the
>> province during the glacial period, and the pattern of recolonization
>> of the province by butterflies specifically and plants and animals in
>> general. It also indicates something of the evolutionary time required
>> for isolated demes to diverge to a consistently different phenotype
>> and (once tested through rearing) genotype. Is this "important"
>> information? The same information could of course be eventually
>> obtained through protein or (once refined) DNA analysis, at great cost
>> for the same detail of geographic pattern, and at some indefinite time
>> in the future. It could be argued of course that there is no need to
>> apply a subspecies name to the phenotypically distinct groups of
>> populations, but there is a definite advantage in being able to use
>> one word to clearly indicate a phenotype that is being discussed.
>
>I don't disagree here at all. However, I should probably let Ken
>Phillip answer this one.  He has an idea about population naming
>that I think is a great compromise. I doubt it will catch on, but it is
>a good idea.  Basically, give whatever population you want an
>appropriate geographical indicator name -- one word will suffice and
>let you know without looking at any label where the bug comes
>from.  I'm sure Ken will correct me if I've screwed up the idea. Sorry
>to put you on the spot, Ken!!
>
>>  Of course, some people
>> name subspecies for the "glory" of it. Personally I name subspecies so
>> that I have a single word which defines a distinctive deme,
>
>Then you *did* understand what I was saying all along!!  Chris, you
>sly devil you!  Since you bombarded me with questions, then I'll
>ask you one, just what do you mean by "distinctive", hmm?  Chris'
>statement represents what I would consider *appropriate* use of
>the subspecies concept.
>
>> Regardless of the motivation for naming a subspecies, a subspecies
>> name results in increased ability to communicate about geographic
>> variation below the species level.
>
>Again, I don't necessarily disagree here, but the geographic
>indicator name idea would seem to be even more useful, instead of
>naming some subspecies after some person.
>
> QUESTION: So what?
>> There are many subjective decisions in systematics, at all taxonomic
>> levels. Why should subjectivity at the subspecies level invalidate
>> that concept, when subjectivity at the species and generic levels does
>> not invalidate those concepts.
>
>Subjectivity will always be a problem with systematics, and there
>will always be disagreements because of it.  Again, my attack was
>perhaps a little too vehement the first time around.  I don't disagree
>with you here.
>
>> "There may be a storm coming as well -- with our increasing ability to
>> be able to pick populations apart at the molecular level, I can see a
>> point where every little molecular difference may be used to say "hey,
>> here's another subspecies (or species)."
> QUESTION: Again, so what? How
>> does this invalidate the subspecies concept any more than the species
>> concept?
>
>Again, I agree, this doesn't invalidate the concept as a whole.  I
>would only disagree with this usage!!
>
>> QUESTION: Why do you interpret the recognition that a species consists
>> of many phenotypically distinct demes, by attaching a subspecies name
>> to each, as being something highly negative?
>
>You, of course, are correct here.  I have had my views slanted by a
>couple of genera.  The two genera I mentioned, Speyeria and
>Euphydryas, have been tremendously split.  I've had many
>discussions with people about western Euphydryas (Checkerspot
>butterflies), and several have mentioned that a number of the
>applied subspecies names seem to be relatively arbitrary.  With as
>much variation as exists *within* populations, it seems quite
>useless (to me)  to have subspecies named whose modal individual
>might be marked slightly differently, but whose variation totally
>encompasses that of another population.  This is just my opinion;
>you may think differently.  And it also bothers me that more
>subspecies are being named without complete knowledge of what
>other subspecies names may actually apply to.
>
> Chris, if names are just supposed to represent geographic
>demes, then I guess I can't disagree with much of anything you've
>said here.  I personally believe, however, that subspecies should
>represent entities that are relatively (hey James, what do you mean
>by "relatively") isolated from one another.  If gene flow is
>widespread among several populations on different mountaintops,
>in different valleys, etc., then I *don't* think calling them different
>subspecies is appropriate, because to me, this is a
>*misrepresentation* of the actual circumstance in nature.  For
>many species, not much is known in many species about vagility
>and gene flow between populations, certainly not to the extent to
>merit calling all sorts of different populations subspecies without
>further study.
>
>> Given some time, we'll probably get around to splitting a bunch of the
>> moths as well, whether they deserve it or not! QUESTION: How could or
>> could not moths "deserve" anything. What criteria do you use for
>> "deserving" splitting?
>
>No offense, Chris, but this was a joke.  Remember those?  I've
>already made it clear that in some cases, splitting may be quite
>useful, wherease in others, it's not.
>
>> SUMMARY: In your entire response there is not a single statement of
>> fact demonstrating that recognition of subspecies is undesirable.
>
>You yourself agreed that some people do throw names out there in
>the case of vanity that are unjustified.  I also hopefully made it
>clearer above why *oversplitting* ("overrecognition") could be very
>undesirable.
>
> Your
>> entire response is a series of opinions.
>
>I didn't say otherwise.
>
>>  I have (above) provided a clear
>> demonstration that recognition of subspecies is a useful biogeographic
>> tool, please provide a clear example where it hinders some aspect of
>> science.
>
>I hopefully have taken care of my rationale to satisfy you, Chris!
>
> Keep keeping me on my toes, Chris!
>
> James
>
>Dr. James K. Adams
>Dept. of Natural Science and Math
>Dalton State College
>213 N. College Drive
>Dalton, GA  30720
>Phone: (706)272-4427; fax: (706)272-2533
>U of Michigan's President James Angell's
>  Secret of Success: "Grow antennae, not horns"
>
>