Demes, meta-populations, subspecies (races) are relatively discontinuous or more-or-less isolated gene pools. Not always completely, but they are to a high enough degree they do not have arbitrary or vague boundaries. This is something Templeton (1998) stresses in his paper. Instead your "Caucasoid", "Negroid", "Mongoloid" are just arbitrary spatial divisions.

I am growing weary of playing pop-up-goes-the-weasel with you.

1. demes/meta-demes need not be relatively discontinuous; show me a definition that requires this.
2. races need not be taxa subspecies (taxa); if they must, explain how nested races are possible; if nested races are not, explain why the literature on race continually discusses them.
3. natural divisions are not necessarily demes. If you think otherwise explain.
4. races were never generally historically conceptualized as being "relatively discontinuous or more-or-less isolated " Thus: http://openpsych.net/forum/showthread.php?tid=226&pid=3504#pid3504 Agree?
5. Templeton's taxa subspecies criterion was made up. If you disagree, explain why he was unable to provide me with a reference which supported his claim and why the reference he provided in his papers didn't. Just search "Templeton" in my paper.
6. If taxa subspecies need to be discontinuous, explain Albrecht, Gelvin, and Miller (2003): "Population structure refers to the geographic arrangement of local populations across the species' range. Population structure can be described in terms of three phenomena: the population continuum, geographic isolates, and zones of secondary intergradation (hybrid zones) (e.g., Mayr and Ashlock, 1991). The population continuum is that part of the species' range where there is continuity of contact among local populations, some of which may be recognized as subspecies if sufficiently differentiated." Previously discussed here: http://openpsych.net/forum/showthread.php?tid=226&pid=3513#pid3513

Please respond to the above.

Tell me why "Caucasoid" is a "natural race", when someone can divide genetic variation in a different way that creates a completely different geographical cluster/spatial division.

Previously discussed (10 pages back): http://openpsych.net/forum/showthread.php?tid=226&pid=3452#pid3452
Also: http://openpsych.net/forum/showthread.php?tid=226&pid=3447#pid3447
You failed to reply. Per biological usage, nature division = genealogical arrangement, not nature made partition (the latter don't exit, since inevitably there is a cross temporal continuum.)

Who is Caucasoid? Where are the boundaries? No proponent of race has ever agreed.

That would be a West Eurasian, a cluster which generally shows up at K=5. Races like populations and demes are relational entities.

At best you cannot argue these "races" are real/natural, but only convenient -- admitting they are arbitrary. For example -

The delineation of all natural divisions is constrained by propinquity of descent, an objective reality. So, along that dimension they are not arbitrary. Given a perfect population continuum, which typically doesn't exist, you could though "arbitrarily" cut out non-overlapping regions of IDB-genomic space. To conceptualize race otherwise would be to conceptualize it inconsistent with historical usage (point 4), no?

Your argument could be we can just split genetic continua so there are "poles" (extremes), despite the boundaries between these "poles" being very imprecise (not natural).

When race is discretely conceptualized, the boundaries would be very precise. There would just be intermediate groups which fell in zones of intergradation. See above.

I understand though it will be hard for you to accept (with a heavily politicized agenda) the truth that your races are nothing more than your own mental constructs.

Is propinquity of descent (ancestry informative molecular relatedness) "nothing more than my own mental construct"? If not than races necessarily aren't either.

I have explained though that race isn't useful -- your arbitrary divisions are only capturing miniscule or trivial amounts of variation.

That would depend on which races we were discussing, no? For example, the average SNP fst difference between North East Asians and West Africans ~ 0.15. The difference between individuals is ~ 1/2*(1-between race), so the ratio of between individual, betwee race to between individual, within race is ~ 0.15/0.425. That's trivial? Given what metric?

You don't explain what use they even have.

Same as "biogeographic ancestry groups". e.g.,
(a) admixture mapping
(b) controlling for population structure in GWAS research
(c) efficacy studies to see if interventions generalize

(I am interested in (a) just with respect to behavioral traits, not boring medical related ones.) This is a dead end for you, since the concept which I justifiably call race is now so widely used.

Here again: http://openpsych.net/forum/showthread.ph...21#pid3521

"I articulated a meaningful sense in which races, so defined, were both real and natural. So you changed the issue to one of whether the race concept and/or its application to humans was "useful". I noted that it was useful for me. And I noted that a number of others employ race or race-like concepts. I also pointed out that the "genetic population", "genetic cluster", and "biographic ancestry group" concepts as often formulated -- ones which clearly are seen as being useful by many -- are equivalent to the general race concept which I was discussing. And I noted that a number of race-critics have pointed out the same, arguing... Recognizing that if you granted that e.g., "biographic ancestry" groups are races "in a phony moustache and glasses" (Silverstein, 2015) you would have to concede the "usefulness" argument, you began to double down on your race-revisionist one..."

...But now you have taken to employing revised race concepts such as race qua metagamodemes to argue against the coherence of my historically grounded race qua natural division concept.

You obviously have nothing. You can't even answer my questions.

You can't even answer my questions.

Here is a list of them. Please answer each.

(1) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3452#pid3452

"Would you at least agree that this latter sense has currency in biology? Dobzhansky summarized the idea nicely -- and even employed the book analogy which you used. In case that it helps, I attached excerpts from his "Evolution Genetics and Mankind", which is available at https://archive.org/

Think the matter over and then let me know if you agree that races, as I define them, form natural divisions in the above sense -- and if this sense has currency"

(2) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3455#pid3455

Do you agree that the entities often called "populations", "clusters", "biogeographic ancestry groups" can, based on historic term usage and insofar as they actually cut out what I call races, legitimately be called "races"?

(3) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3462#pid3462

I don't think that Garn equated races with demes -- but I will have to read more of his articles to be sure. That is, I imagine that his races were like Dobzhansky's where while all races are "Mendelian populations", not all Mendelian populations -- specifically demes which were not genetically "distinct" -- are races. Thus, he notes that his races share a fraction of genes in common. Strictly speaking demes do not need to. Would not you agree?

(4) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3480#pid3480

You are making one of two arguments here:
(1) "race concept isn't useful because there is more variation in populations than between them" which precludes the accurate classification of humans into racial (i.e., genealogy-based) groups.
(2) "race concept isn't useful because there is more variation in populations than between them" which means that differences between said-human groups are of no importance.

Could you clarify which one? ... Now, before I discuss the problem with argument (2) above, I would like you to clarify whether you mean (1, or 2). If (2), could you clarify the logic? Specifically, how would you quantify "importance" in terms of average racial difference?

(5) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3481#pid3481

Here, I see two arguments:
(a) race typologists in the early 20th century (generally) argued that the variation between races was non-trivial/major.
(b) race theorists prior to Boas (generally) argued that the variation between races was non-trivial/major.

Would you agree that monogenists such as Prichard argued that the variation was relatively unimportant -- since it was of the intraspecific type?

A problem with both claims about the bimetric typologists is that "non-trivial/major"/"trivial/minor" is a somewhat subjective estimate; as such, it's difficult to put claims on a common metricand to evaluate if they overestimated differences.

So how would you propose to evaluate early 20th century qualitative claims? I would think that we would just look at their quantitative data. Isn't it reasonable to assume that they thought that differences were no larger than their reported data showed? If so, since they present the data with means and standard deviations we can just look at that. If it shows differences no larger than what is now known, we can conclude that they though that differences were no larger or significant than they quantitatively are. If so, then I would just have to show that their metrical data was not greatly off to show that their evaluations were not. Do you agree?

(6) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3489#pid3489

Naturally enough, I explicitly formulated the concept such to be consistent with divergence due to isolation by distance (e.g., page 41) -- which is to say a continuum due to primary intergradation. This is what distinguishes it from e.g., Shiao's "clinal class" one (see page 64).

Now you quote:

"Therefore, there is no reason to assume that major genetic discontinuities exist between different continents or races."
http://genome.cshlp.org/content/14/9/1679.full

You know that I know the history of this debate at least as well as you do, so I don't see why you would cite the above. Shiao (2014) addressed this in his defense of his clinal class concept. But of course, you are right, my stance is:

(a) one could "arbitrarily divide" a "genetic continuum" into races.

Of course, I anticipated objections to this position and discussed them lengthy. Which parts of my discussions didn't you find convincing? Do you want me to refer you to the specific sections or to elaborate on specific points?

(7) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3489#pid3489

Pearson's comment suggests that he was overestimating the relative magnitude of between group differences -- though it is not clear as it's not clear if Pearson was discussing multivariate distance (his coefficient of likeness, which was a primitive form of Mahalanobis distance) or average differences. Whatever the case, the very occurrence of this discussion calls your claim of a general view of homogenous races into question. And once you admit that variability was recognized, you are forced to double down on your claim that biometric typologists felt that variability within populations arose from the admixture of distinct homogenous groups. Yet it is clear that they didn't believe in originally distinct homogenous races. Thus Pearson notes....

If they diverged from a common stock in continuous degree and now contain enough variability such that a " Coefficient of Racial Likeness" is needed when and how were they ever homogenous groups in the sense you mean?

(8) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3491#pid3491

To preclude confusion, let's distinguish issues:

(1) whether or not the natural division concept of race is coherent
(2) whether or not such and such characters are reliable indexes of evolutionary relationship (and thus reliable indexes of racial membership)

Recall this discussion on reliability
http://openpsych.net/forum/showthread.ph...47#pid3447

Could you clarify which point you wish to discuss in relation to the forensic data?

(9) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3497#pid3497

As held by whom? Buffon, Blumenbach, Esper, Prichard, Darwin, Quatrefages ...? Specifically, list the many early (hence "traditional") proponents of race (as divisions of a species) who maintained that these divisions were -- moreover were definitionally so -- marked by "abrupt change" or "sharp discontinuity". I do not want to hear Naomi Zack's bullshit; I want specific "traditional" references.

(10) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3509#pid3509

But I like to settle one point at a time. When you concede that races per se were not typically thought of as deeply discontinuous and were often recognized to be continuous I will revisit the question of original -- "in situ" -- homogeneity.

[Do you concede this?]

(11) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3513#pid3513

Krom: Grover Krantz was a proponent of race. Unlike you he wasn't trying to redefine the concept.

How possibly are his "micro-races" qua micro geographical populations more authentic than my natural division ones?

(12) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3518#pid3518

Krom: but the fact most biologists do not regard categories like "Caucasoid", or "Negroid" to be useful.

Could you point us to the surveys which show that most biologists do not consider such and such race concepts to apply meaningfully to modern humans?

(13) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3566#pid3566

Do you agree that when an author claims that there are "large" differences between such and such races this does not entail that her concept of race necessitates such differences? Yes or no?

(14) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3574#pid3574

Would you honestly argue that this wasn't revisionist? (Please answer.) That early race concepts proposed treeness in Templeton's sense of having little admixture, great isolation, and significant discontinuities (Didn't we discuss this?) or that early concepts required high between group genetic variation -- when the concept of "gene" was not developed until the late 19th to early 20th century. Or that race was always identified with subspecies, when the race concept preceded that latter and given that authors often drew the distinction between subspecies in the taxonomic sense and race. Or that early subspecies concepts required the types of differences now seen as sufficiently taxonomically significant -- actually there are no formal criteria; and the only generally accepted rule of thumb is the 75% one, by which major human races qualify as taxa subspecies --for BSC subspecies recognition.

(15) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3585#pid3585

To sum up, if you wish to argue that race is only a useful classification if all races in a given species at a given time are given names then you must either (a) maintain that races must be taxa or (b) contend that same holds for all other like classifications e.g., "deme", "(spatial) population", "morph", "biotype", "stock", "strain", "cluster", "form", "ecotype", "groups" (i.e., assemblages of closely related taxa), "semispecies", and "super-species". Which will it be? You will have to adopt either an absurd position (b) or a revisionist one (a).

2. Regarding races and levels of analysis, which of the following would you like to maintain were not infrequently recognized as races (in the sense which I mean)?


(16) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3595#pid3595

Esper is saying that subspecies/races are not the same as inconstant varieties ("plane ab his sunt separandae") because like species, and unlike inconstant varieties, they reproduce their separate type ("Ad procreandam sobolem eamque ipsis aequalem aptae"). But, he notes, they are also not like species, because they originated from species ("Originem ex speciebus duxisse"), similar to how inconstant varieties do ("caussa qua fuere mutatae eadem manente"), which is obvious because they lack essential or species-like differences ("perfectus in iis declarat partium essentialium similitudo"). That is, according to Esper, different subspecies/races, while distinct ("Characteríbus autem pariter sunt distinctae") are essentially the same and thus unlike species.

Now, perhaps you can explain how you get "subspecies require "perfect similarity" in certain traits" from " Originem ex speciebus duxisse, perfectus in iis declarat partium essentialium similitudo". And while you are at it, perhaps you can explain how Esper's statement that subspecies are while different essentially the same fits with your narrative.

(17) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3601#pid3601

For example, North and South America form two massive genogamodemes, but surely not two biogeographic ancestry groups or natural divisions. Thus, what I call race -- and what was often called race -- is not redundant with what you call deme.

So that we can move forward, can you either concur or disagree?

(18) Here: http://openpsych.net/forum/showthread.php?tid=226&pid=3603#pid3603

If my race concept is not useful, then why is it so often used, just under different names e.g., "biogeographic ancestry groups", "genetic clusters", etc?



Thanks.

P.S. from my last post:

2. races need not be taxa subspecies (taxa); [Do you agree?] if they must, explain how nested races are possible; if nested races are not, explain why the literature on race continually discusses them.

3. natural divisions are not necessarily demes. [Do you agree?] If you think otherwise explain.

4. races were never generally historically conceptualized as being "relatively discontinuous or more-or-less isolated " Thus: http://openpsych.net/forum/showthread.ph...04#pid3504 Agree?

5. Templeton's taxa subspecies criterion was made up. [Do you agree?] If you disagree, explain why he was unable to provide me with a reference which supported his claim and why the reference he provided in his papers didn't. Just search "Templeton" in my paper.

6. If taxa subspecies need to be discontinuous, explain Albrecht, Gelvin, and Miller (2003): "Population structure refers to the geographic arrangement of local populations across the species' range. Population structure can be described in terms of three phenomena: the population continuum, geographic isolates, and zones of secondary intergradation (hybrid zones) (e.g., Mayr and Ashlock, 1991). The population continuum is that part of the species' range where there is continuity of contact among local populations, some of which may be recognized as subspecies if sufficiently differentiated." Previously discussed here: http://openpsych.net/forum/showthread.ph...13#pid3513

...

The delineation of all natural divisions is constrained by propinquity of descent, an objective reality. So, along that dimension they are not arbitrary. Given a perfect population continuum, which typically doesn't exist, you could though "arbitrarily" cut out non-overlapping regions of IDB-genomic space. To conceptualize race otherwise would be to conceptualize it inconsistent with historical usage (point 4), no?

....

Is propinquity of descent (ancestry informative molecular relatedness) "nothing more than my own mental construct"? If not than races necessarily aren't either [, no?].

....

That would depend on which races we were discussing, no? For example, the average SNP fst difference between North East Asians and West Africans ~ 0.15. The difference between individuals is ~ 1/2*(1-between race), so the ratio of between individual, betwee race to between individual, within race is ~ 0.15/0.425. That's trivial? Given what metric?

The *ideal* deme is 100% a mating isolate or gene pool (this means 100% of x individuals choose sexual mates within x, hence random mating: panmixia). This is rarely found, so demes by definition are -more or less isolated-, meaning sharply (but not always wholly) discontinuous local breeding populations where out-group mating (i.e. gene flow) is minimal

How do you deduce "sharply" from "more or less isolated"?

Would you agree that you can cut demes out of a breeding continuum? Or do you imagine that you need sharp barriers separating any two demes?

Please answer this.

[Added:

If demes exhibit "sharp" differences owing to "spatial distinctiveness" then natural division races do also, since the latter are located in distinct (i.e., non-overlapping) regions of genomic space.]

redundant.

Wright (1978) wrote the following about Fst -

“The range 0 to 0.05 may be considered as indicating little genetic differentiation.
The range 0.05 to 0.15 indicates moderate genetic differentiation.
The range 0.15 to 0.25 indicates great genetic differentiation.
Values of FST above 0.25 indicate very great genetic differentiation.”

The very low stats (which fall under Wright's "miniscule") are not discussed in your book.

I'll put my conclusion first, since it's obvious that you don't bother reading through:

So, for continental groups, using Mt/YDNA you get great differences. Using microsatellites and common methods such as Nei's or Weir and Cockerham's Fst, you get lower-end moderate ones. Using Jost's D or other similar measures which correct for Hs, you would get, by this scale, great differences. Using SNPs, the more relevant loci, you get upper-end moderate ones. Only by cherry picking -- AMOVA applied to microsatellites -- do you get the results you desire. How about that? And of course those results are misleading because (a) Wright discussed Fst, not AMOVA -- thus rendering problematic the use of Wright's Fst based scale, (b) using typical statistical scales, the 3.8% is out of < 30% max -- thus rendering problematic a straightforward percent comparison -- we actually are not dealing with percents, and (c) the whole genetic differentiation comparison is misleading, in context to discussions of quantitative trait differences, because between group differences are out of inter+intra individual ones.

As for phenotypic differences, it depends on which you look at. Some are larger than what you would expect given the SNP fst differences, indicating divergent selection e.g., Guo et al. (2014). Variation and signatures of selection on the human face. Journal of human evolution, 75, 143-152.

Some are about the same size e.g., craniometric and dental, indicating neutral variation. Some are smaller, indicating stabilizing selection.

My point would be, unless I am missing something, that the default prediction for quantitative trait differences would be ~ 2 x SNP Fst. And this is large by social scientific standards. Now I asked some, who had written on the topic, such as Henry Harpending, and they thought, from what I recall, that the above didn't sound too far off. But if you can find a flaw in the logic, let me know, I would appreciate it.

...

The longer version is:

In fact, I do discuss this issue extensively and I cite that very rule of thumb. I show both microsatellite and SNP differences -- see picks below. I discuss how differentiation values depend on both the method and especially the loci. I note that fst values are constrained by heterozygosity, thus the Nei's Fst of 0.055 found by Rosenberg et al. (not to be confused with AMOVA) is out of a maximum of 0.28. When you correct for heterozygosity using, for example Jost's D, you get a value of 0.20 (crudely 0.055/0.28). This is closer to the SNP difference, which is higher because SNPs have lower heterozygosity (around 0.30, correcting for Hs ~ 0.17). I noted:

"In short, species with high within population diversity will necessarily show low between population Fst and Fst analog values, regardless of the actual between population diversity as indexed by the number of shared alleles. (Readers are referred to Jakobsson, Edge, and Rosenberg (2013) for a discussion of this matter with regards to human populations.)...Thus, the low microsatellite Fst value between major human races is about what one would expect to find were one dealing with the subspecies of a species which had a heterozygosity value similar to that of the human species...."

And later:

"Whitlock (2008) explains the measure Qst: The calculation of QST for a trait requires two quantities: the additive genetic variance of the trait within a population (V A, within) and the genetic variance among populations (V G, among). For diploids, QST is calculated as:

Qst = V G, among / (V G, among + 2V A, within)

For haploids, the same equation applies, but without the '2' in the denominator.[That '2' for the diploid case comes from the fact that the quantitative genetic variance among populations is proportional to two times FST (Wright, 1951).]

What is particularly relevant to the present discussion is Whitlock's last statement, since we are interested in predicting quantitative genetic variance from Fst, not comparing Qst to Fst. Amongst diploid populations, the predicted quantitative genetic trait variance is equal to 2Fst/(1 + Fst) (Leinonen et al., 2013). The 2 in the equation comes from the fact that roughly half of the genetic variation within diploid populations is within individuals....

Getting back to the main point, if we wish to estimate expected quantitative genetic trait variation it is often advised to avoid using low mutation rate genetic markers such as microsatellites, which, as discussed, have high Hs values and thus necessarily exhibit low Fst values. It is often advised instead to use SNPs, both because these markers do not tend to have very high Hs values and because SNP variation codes for typical quantitative trait variation (Edelaar and Björklund, 2011). Another way to look at this is to consider that the magnitude of (fixation index estimated) genetic differentiation varies by the class of loci analyzed, with part of this variation being attributable to loci variation in Hs (Jakobsson et al., 2013); for example, for humans, continental microsatellite, SNP, and mtDNA Fst values are typically around, respectively, 0.05, 0.12, and 0.20. Were one to try to infer the magnitude of genetically conditioned phenotypic variation from typical indices of fixation (e.g., Fst values), it would make sense to use the class of loci that most likely underpins the relevant trait variation. For example, since variation in single-nucleotide polymorphisms (SNPs) explains variation in many interesting polygenetic traits such as height and intelligence (for example: Yang et al., 2010; Davies et al., 2011), it would make more sense to attempt to infer magnitudes of genetic differentiation in these traits from SNP Fst values than from microsatellite or mtDNA ones." Blah blah blah ...

Would you agree that you can cut demes out of a breeding continuum? Or do you imagine that you need sharp barriers separating any two demes?

I keep encountering papers which say something to the effect of:

"In general, the variation in estimates of pairwise genetic distances explained by these effective distances is compared with that explained by geographic distances alone (that is, IBD). The latter is regarded as the most simple landscape genetic pattern that would be obtained even if there were no landscape effects and migration was thus only constrained by distance between demes (Spear et al., 2005; Balkenhol et al., 2009; Jenkins et al., 2010). This notion may have originated from spatially explicit simulation studies of IBD patterns, in which demes or individuals are usually placed in regular lattices throughout homogeneous spaces (Guillot et al., 2009; Epperson et al., 2010). Indeed, distance-constrained migration in such
models produces IBD patterns that are not influenced by any landscape elements. However, a heterogeneous landscape will not only affect migration probabilities between demes, but also the spatial arrangement of demes (that is, deme topology)"
http://www.nature.com/hdy/journal/v114/n1/pdf/hdy201462a.pdf

The authors seem to have no problem with discussing demes in context to isolation by distance i.e., a continuum... but, I imagine that you are more familiar with the literature, so I await to hear your reply.

My point was that if you are going to say that demes picked out of a continuum are necessarily sharply differentiated, you have to grant the same for races (as I define them).

If you don't wish to do the latter, you either have to say:
(a) demes can't be picked out of a continuum (interesting claim for which I would like to see support)
or
(b) demes need not be sharply different (as I originally argued)

Which will it be?

Only by cherry picking -- AMOVA applied to microsatellites -- do you get the results you desire. How about that? And of course those results are misleading because (a) Wright discussed Fst, not AMOVA -- thus rendering problematic the use of Wright's Fst based scale

I actually included a table with all published continental level micro Fst values I could find. Attached.

Are you aware Rosenberg et al (2002, 2005) clusters are said to only capture 1.53% inter-population genetic variation. >98% is captured by IBD.

Now, Rosenberg (2005) said:

"When an additional binary variable B is added—equaling one if an ocean, the Himalayas, or the Sahara must be crossed to travel between two populations, and zero otherwise—R2 increases to 0.729. The regression equation is Fst = 0.0032 + 0.0049D + 0.0153B, where D is distance in thousands of kilometers....The effect of a barrier is to add 0.0153 to Fst beyond the value predicted by geographic distance alone."

The 0.015 is just the effect of their crude dichotomously coded barriers; this is not the percentage of the total variability between continental races (K=5). For that you have to turn back to Rosenberg (2002). Rosenberg (2002) gives an Amova of 4.3 between regions (not 3.6 as you said earlier!) for K=5 (Blumenbach races! Remember, Linnaeus had inconstant varieties!). But what does this 4.3% actually mean? For context, try:

.....................

Meirmans, P. G., & Hedrick, P. W. (2011). Assessing population structure: FST and related measures. Molecular Ecology Resources, 11(1), 5-18.

"For biallelic markers, this makes sure that FST is bounded between zero and one, with zero representing no differentiation and one representing fixation of different alleles within populations. For multiallelic markers, however, the maximum possible value is not necessarily equal to one, but is instead determined by the amount of within-population diversity (Charlesworth 1998; Hedrick 1999). The reason for this can be best understood by looking at GST, which is defined as (Nei 1987)....For highly variable loci, this can lead to a very small possible range of GST values. To illustrate this relationship, Fig. 1 gives the joint values of FST and HS found in the past 4 years in Molecular Ecology (expanded from Heller & Siegismund 2009; see also Table S1, Supporting information). Notice that the observed range of FST is always less than HS and that the range of FST becomes very small when HS is large. For example when HS = 0.9, a value that is commonly encountered for microsatellite markers, the maximum possible value of FST is 0.1. Such a value of FST is generally interpreted as representing a rather weak population structure. However, here it represents the case with maximum differentiation among the populations, meaning that the populations do not share any alleles at all."

But see also here: Verity, R., & Nichols, R. A. (2014). What is genetic differentiation, and how should we measure it—GST, D, neither or both?. Molecular ecology, 23(17), 4216-4225.

"By tracing GST to its origin – the parameter FST, and further to the inbreeding coefficient F upon which FST was built – we can identify the root cause of some of the disagreement in the literature around the measurement of population differentiation. We have found that there are two overlapping views regarding the definition of the probability of identity by descent, which in turn have rubbed off on our definitions of FST, leading to mutation dependent and mutation-independent versions. The criticism that GST is constrained is misplaced – at least if the task at hand is to estimate the mutation-dependent version of FST. If we wish to capture other aspects of the population history, such as the mutation independent version of FST, when the mutation rate is relatively high, then we will need to supplement GST with other measures that capture a different aspect of evolution. No single statistic can be informative about both parameters in this situation, as it is mathematically impossible for a single dimension to fully represent two.... Thus no statistic is differentiation, but some statistics can be used to infer differentiation. We find that mutation-independent FST is a sensible quantity to use as our definition of differentiation, although the general arguments made above are equally valid when applied to alternative definitions... Under the same model with a high mutation rate we found that GST is insufficient on its own to jointly estimate the true level of differentiation and mutation (Figure 5). Supplementing GST with either G’ST or D solved this problem, providing a distinct source of information that can be used to pull apart the confounded signals."

And here: Should I use FST, G’ST or D?

"Over the past few decades researchers have increasingly used microsatellites, due to their high level of variability and the relative ease of development and scoring in non-model systems. However, now that next-generation sequencing is getting more affordable, sequence-based markers can be assessed throughout the genome (e.g. using RAD sequencing). As we move back towards such low-mutation-rate markers as SNPs, FST becomes easier to assess reliably. On the other hand, FST and other current methods are all designed to assess one or a few markers at a time, and genomic approaches just apply these methods thousands or tens of thousands of times for markers throughout the genome. One can look for outliers, calculate means, etc., without really taking full advantage of the data. For instance, I have seen bi-modal or skewed distributions of FST and other summary statistics; clearly means and standard deviations can be misleading in these cases. My hope is that new methods for assessing divergence will focus not on individual loci but on many markers throughout the genome....
The theoretical maximum of FST = 1 can only be reached if each subpopulation is fixed for a single unique allele. If there is variability within any subpopulation, the maximum FST is (1 – HeS). Unfortunately, this limit to the maximum FST is often overlooked. The maximum value for FST is the smaller the more variable a marker is, and the effect can be especially dramatic for microsatellites, which often exhibit high HeS (over 0.9, in which case the maximum FST is only 0.1). In the extreme (yet possible) scenario of two subpopulations completely divergent (i.e., not sharing a single allele), but both with HeS approaching 1 (i.e., all individuals are expected to be heterozygous because of high allelic diversity), FST becomes meaningless, as its theoretical maximum is then 0 (see Fig. 1 in Jost 2008 for a graphical representation)"

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I'm not 100% sure how Amova -- which is a F/Gst Analogue -- works, so I will just use F/Gst values, which were about 0.05 (rounding down). The clearest interpretation would be: (a) the between regional diversity accounts for 5% of the total diversity, (b) given an upper limit of 28% for these loci. (c) This is equivalent to a 20% mutation independent differentiation (measured in allele sharing) (e.g., Jost's D). Now claim (a) is fine so long as you understand what F/Gst is measuring and recognize its dependency on mutation rates. But (a) is not fine if you wish to make a global claim about between population differentiation and base it on high mutation rate loci. On this point, It is notable the Wright (1969) developed Fst for biallelic markers, which as Meirmans and Hedrick note actually have a range of 0 to 100. So it's not clear how well Wright's scale transfers to multiallelic markers, which are constrained by Hs. On this point, Nolan Kane notes (correctly in my opinion):

"For many situations, certainly, this can be quite problematic – for microsatellites with high heterozygosity, maximum GST is often 0.1-0.2! Clearly, in these cases Wright’s (1978) guidelines are entirely misleading, when he states that values ranging from 0-0.05 indicate “little” genetic differentiation; 0.05-0.15 is “moderate”, 0.15-0.25 is “great”, etc. This is only plausible for biallelic cases, and in other situations we cannot rely on such simple rules of thumb. (Should I use FST, G’ST or D?)"

So, to summarize, yes, Human CT microsatellite F/Gst is low to moderate. But this is expected given the high heterozygosity of the markers. (Indeed, my regression line for subspecies Hs by G/Fst, showed that the human micro G/Fst was what one would expect for a subspeciated species with the same level of Hs!) More generally, since mutation independent measures of differentiation give estimates which substantially diverge from those given by mutation dependent ones, it's difficult to interpret the situation in terms of "general" differentiation. Also, since Wright's rule of thumb was based on biallelic Fst (which has a practical range of 0 to 100), it's not clear to what extent it is applicable to multiallelic Gst, specifically when using high mutation rate microsat.

Luckily, the above is somewhat tangential to the immediate discussion, which concerns expected quantitative differences. I say luckily, since the issue above is too complex for me to easily convince you of the point, given your stubbornness on the topic.. I say that it's tangential, because unless I am mistaken, you are arguing that as (micro) Fst is low, typical quantitative differences must also be so. Conveniently for me, rules of thumb for making these inferences have been outlined. (references cited in my paper.) And they note that when doing so, when assessing the expected trait differences owing to neutral divergence (the default magnitude of differences), one should use low mutation rate markers of the type which likely underlie the genetic structure of the trait, such as SNPs, for which, amongst humans, there happen to be moderate to large differences, depending on the groups discussed. I elaborated on this in my paper -- section 4 -- and so I won't rehash all of the points made. What I said, though, definitively refutes these types of silly arguments which you are reiterating.

Your move.

You talk about cherry picking but you don't calculate a mean from all these studies. I take them all into account and there is more than above. A lot more genetic markers have very low values of under 5%. Obviously those traits like skin colour and limb have to be excluded because they are not neutral genetic/phenotypic markers.

I replied to an earlier point above.

Look, a Fst value of 10 to 15% is large and this is what you find with the SNP data, both in 1000 Genomes and HapMap. (Barbujani is fond of citing the unweighted 1000 Genome Fst values, reported in the original 1000 genomes report.*) For making inferences about quantitative differences owing to neutral selection, this is the preferred value:

"One valid solution is to estimate FST from neutral markers with mutation rates comparable to those of the loci underlying quantitative traits (e.g. SNPs). QST can also be compared to ΦST (PhiST) of amova, as long as the genetic distance among allelic variants used to estimate ΦST reflects evolutionary history: in that case, neutral divergence is independent of mutation rate. In contrast to their common usage in comparisons of QST and FST, microsatellites typically have high mutation rates and do not evolve according to a simple evolutionary model, so are best avoided in QST–FST comparisons."

Plugging in the SNP Fst differences into the equation for expected quantitative differences, gives you large differences. I discussed this in my paper. Now for phenotypic Fst (i.e., Qst values) you give:

(1) Pelvic shape (males) 2.6% Pelvic shape (females) 3.3% --
(2) Craniometrics is 14.6% as calculated by Relethford (1994, 2002) -- large
(3) Limb ratios are around 38% Fst. -- very great
(4) Skin colour is 87.9% Fst (Relethford, 2002) -- very great

All but (1) represent large to differences, as judged from the perspective of the social science. (See my table 4.12. Attached below.) And this is neglecting the fact that Qst is calculated as:

Qst = V G, among / (V G, among + 2V A, within),

with a 2 in the formula to account for diploidy, which means that expected quantitative trait differences:

2Fst/(1 + Fst) (Leinonen et al., 2013)

These are large numbers. (Oh yes, and I emailed Barbujani about this, but he didn't care to reply.)

Maybe you are just thinking of a different scale. But since we are talking about behavioral and medically relevant differences, the scale I am using is arguably the correct one, since this is the one people use in the biomedical and social sciences. For example, the SES variance between Whites and Blacks in the US is around 7% (medium effect size differences, measure depending). No one says, "Oh these don't exist, so there's no reason to study sociologically delineated race". Why? Because these magnitudes of differences are not typically interpreted as being irrelevant. Now maybe you disagree. Maybe you feel that sociological race classifications are empirically meaningless because most the of variance is within populations. Do you? Whatever the case, for better or worse I am following the common evaluative standards, which leads me to the expectation that there are "important" behavioral differences, where "importance" is evaluated according to common standards for judging inter-individual differences.

Now, that returns us back to the issue of "the" magnitude of genetic differences. There is of course no "the" because estimates vary according to the races under question, the loci analysed, and the method used. Since I am not particularly interested in continental level divisions -- for various reasons, some empirical some ideological -- I do not fixate on these differences. Rather, I look at differences between e.g., Europeans, West Africans, and Amerindians, which given my construction of the race concept and consistent with the biogeographical ancestry one, I can treat as three separate racial groups.

But for some reason you are interested in continental-level differences. In interpreting these, as I noted, you have to keep in mind mutation rates, both if you want to make interspecies comparisons or interindividual, between race ones.

serum proteins and red blood cell enzymes 9% (Latter, 1980)
X chromosome STR's Ramachandran et al. (2004) 4.9%
Alu insertions Romualdi et al. (2002) 8.9%
642,690 atDNA SNP's Li et al. (2008) 9.0%
HLA 7.0% Meyer et al. (2006)

Now you said, "you don't calculate a mean from all of these studies". That is correct and for good reason. It makes no theoretical sense to try to average e.g., short tandem repeats (microsatellites), which are picked specifically because of their high mutation rates, and e.g., SNPs, which have relatively low rates to try to get a "true differentiation rate", without even a clear idea about what you mean by "differentiation", or to combine values made using different estimators, different methods of loci aggregation, different methods of handling rare alleles, ect. If Barbujani in fact did this, I would have to guess that he was being intentfully misleading -- and if it was just you, that you don't have a good grasp of this issue.

In my paper, I believe I said: "[F]or example, for humans, continental microsatellite, SNP, and mtDNA Fst values are typically around, respectively, 0.05, 0.12, and 0.20...The median continental race SNP Fst value is said to be around 0.12 (Li et al., 2008; Campbell and Tishkoff, 2008; Elhaik, 2012; Bhatia, Patterson, Sankararaman, and Price, 2013)".

This sounds about right, though I guess we could conduct a more thorough review.

The point here though is that discussing average "differentiation" values is meaningless:

(a) If we wish to make an interspecies comparison to assess the magnitude of e.g., continental human differentiation relative to that between the formally recognized races of other species, then we would have to pick a specific class of loci and compare. Both Templeton (1998) and I did this and found that humans fell around the 15th percentile (low average).

(b) If we wish to use Wright's metric to judge the magnitude of continental-level differentiation then we would need to again use appropriate loci -- presumably biallelic, but we could check what he used.

(c) If we wish to determine the expected quantitative trait difference between e.g., continental races, then again we need to use the appropriate loci e.g., SNPs for common morphological/behavioral traits; perhaps others for biomedical ones. Presumably not high mutation ones such as STRs.

*In "Genes in space Selection, association and variation in spatially structured populations", the authors notes:

"On these scales, the 1000 Genomes Project Consortium (2010) reported pairwise FST of 0.07 between European and West African populations, 0.08 between East Asians and West Africans, and 0.05 between East Asians and Europeans. These numbers are unweighted averages over all sites, and are therefore lower (empirically, by a factor of about 2) then the corresponding weighted average"

“It is likely that each breeding population will prove to be genetically unique, so that all will be racially distinct in Dobzhansky’s terms. But this is not the general use of the concept of race in biology, and the concept has not in the past been associated with this theory of human diversity.” (Livingstone, 1962)

“There are undoubtedly no two genetically identical populations in the world; this has nothing to do directly with the validity of race as a taxonomic device. Unless we have defined exactly what we mean by this… differences between populations are population differences, nothing more.” (Hiernaux, 1963

Ok, but as said, natural division races are not equivalent to genogamodemes in the sense of breeding populations which differ in some genes. Do you grant this point or shall we keep playing pop up goes the weasel? For example social classes and religious groups would often be the latter, but would not typically constitute natural division races (unless they were sufficiently linebred). As discussed prior, I am not sure that Dobzhansky’s races were just genogamodemes -- that is I'm not sure that all of his genetically different Mendelian populations were races. Whatever the case this genogamodemes concept offers a nice contrast with the natural division one, which requires individual to be more overall related to other members of their groups than to members of other groups. And yes I agree that many genogamodemes would not make for good races.

Demes or meta-populations (groups of demes) with very few exceptions are not ethnic groups.

When I google "deme" and "degree of isolation, I get e.g.,:

"Instead, the population of the species is subdivided into a number of demes, which are isolated from one another to varying degrees, either by unfavorable habitat or simply by distance. Given some degree of isolation, it becomes highly probable, indeed inevitable, that some genetic divergence among the demes will occur". (Merrell, D. Ecological Genetics)"

"No extra operations are introduced as a results of distributing the populations into overlapping demes. The only factors that vary in these experiments are degree of isolation..." (Ebeling et al. "Parallel Problem Solving from Nature)

There is no clear degree of isolation, demes can overlap, demes can fall on a continuum, demes per se don't necessarily have genetic differences.

So I find this odd: https://books.google.com/books?id=yx8yBwAAQBAJ&pg=PA142&lpg=PA142&dq=%22ethnic+groups%22+%22demes%22&source=bl&ots=msbKT6uZEt&sig=Nh_6Ln5n6iij2AoOZb45PakGdmM&hl=en&sa=X&ved=0CB4Q6AEwAGoVChMIuqq0_e_txwIVR8yACh16SQtj#v=onepage&q=%22ethnic%20groups%22%20%22demes%22&f=false

Nordgren's demes look just like my races: "Demes are groups of individuals who are more similar to each other than to any other individuals". And they look quite unlike the demes of others e.g., "Given some degree of isolation, it becomes highly probable, indeed inevitable, that some genetic divergence among the demes will occur". If you can find a meta-review of the "deme" notion send it along, because it looks like many are using the term to reference very different entities.

Anyways, this is what I like to focus on concepts not terms. What was a Krom-demes, again? If you wish to say that demes need a very high degree of reproductive isolation ("more or less" is pretty ambiguous), how do you justify a cut off and what do you call less isolated groups ... incipient demes?

Ethnic groups (e.g. "Germans") are mostly arbitrary spatial constructs like races, e.g. "Caucasoid".

I don't imagine that most would define a German strictly spatially e.g., such that a "German" could never exist outside of "Germany". With the amount of immigration, I also don't imagine that "Germans" in the political sense even constitute and "ethnic" group in the anthropological sense of groups of individual with a shared identity, history, heritage... Though, there are ethnic Germans and some e.g., ethnic Germans belong to a different natural division race than some political/ethnic Chinese. Caucasoid is easier since this isn't an ethnic identity or a political construct. One can stipulate that Caucasoids refer to members of the West Eurasian biogeographic ancestry group in which case they would constitute a genetic construct (biological race, my sense). They don't form, as typically used, an obvious spatial construct (spatial population) since "Caucasoid" usually refers to Whites in Australia but not Black African in Europe. Would they be a social construct -- well even if they formed a taxa subspecies they might be, so this is a pretty non-informative category. Maybe a cluster-morph?

Despite the standard scientific/textbook definition of a (gamo)deme is a more-or-less isolated breeding group/mating circle.

Why not just type "gamodeme"/"deme" and search Google books. Its not hard.

http://www.merriam-webster.com/dictionary/gamodeme

You missed the point as usual. (I wasn't discussing degree of isolation here, but location on a continuum.)

(1) Demes can be cut from an isolation by distance breeding continuum. True or false?
(2) If you want to say that these demes which fall on a breeding continuum are "sharply differentiated", then you must also say that my discretely conceptualized (non-overlapping) races cut from a continuum are likewise "sharply different".

I am fine with that. Just acknowledge the point. It's odd that you can't see the symmetry.

Ignoring genetic differentiation for a moment, races are intergenerational aggregates of demes.

Are you saying that Krom-races are these or that Chuck-races are? That is, who's concept are we discussing now? Or are you arguing that this is what "race" generally should mean (general vesus narrow concepts), given historical usage? If the latter, what's your justification for this particular broad concept? And how isn't this definition "revisionist", given that the "deme" concept is relatively new? And isn't this thread concerned with my race concept? Or are you now arguing that mine mismatches with what race, broadly understood, really means (according to Krom)? Clarify please!

As Templeton explains, races must be "geographically circumscribed". This doesn't just mean arbitrarily marking boundaries across space as you are doing for "Caucasoid", "Negroid", "Mongoloid". Your races are not "natural" and don't exist anywhere but in your mind.

As Templeton explains... Templeton couldn't even distinguish between a morphological rule of thumb (the 75% rule) and an imaginary genetic rule. Moreover, he couldn't recognize the difference between taxa subspecies and races. As I asked prior, if races = taxa subspecies, how can there be nested races? But now -- according to you, at least -- Templeton's taxa subspecies -- he doesn't discuss race in a more general sense (e.g., "race" versus "geographic race") -- must be (prescriptively) "geographically circumscribed". If this is generally the case, why, I ask, do formally recognized races of plants and animals which are transported around the world to zoos in heterogeneous locations retain their trinomen? Whatever the case, the modern taxa subspecies / "geographic race" concept is mostly an appendage of Mayr's biological species concept. So we can simply turn to Mayr for guidance on this murky issue:

"When dealing with human races we must think of them as the inhabitants of the geographical regions in which they had originated... A human race consists of the descendants of a once isolated geographical population primarily adapted for the environment conditions of their original home country... But, as is illustrated by the success of Europeans and Africans and Asians in all parts of the world, any race is capable of living anywhere." (Mayr, 2002 "The biology of Race and the Concept of Equality")

Mayr's geographic races -- and therefore BSC ones -- hardly unconditionally necessitate geographical circumscription (they just have to had descended from geographically circumscribed populations). Which is why non-human races in zoos spread around the world are treated as they are.

But of course maybe Templeton's taxa subspecies based on Templeton's cohesion species concept are different. If so then, by his idiosyncratic concept there perhaps could be no human taxa subspecies. But who cares?

I'm finishing this dispute, but here is my summary of what I call elsewhere (following Hochman) "weak racial population naturalism":

The fact that I was able to respond to all of your numerous critiques and questions (some 20+ pages of them) while you were unable to return the favor testifies either to the coherence of my defense or the incoherence of your critique or both. But you add more, so let me address that briefly.

If Glasgow (2003) is right that we must stay fairly near to the original meaning, then your supposed "standard taxonomic definition" can be ruled out as a general concept. Also, if he is right, then Hochman's (2013) "strong races" can also be. Historically, intraspecific race just described (genealogically understood) contant varieties, the differences between which could be fairly trivial. More generally, race described lineage, specific or intraspecific. For example, naturalist Georg Forster noted in 1786:

We have borrowed <the term> [race] from the French; it seems very closely related to <the words> racine and radix and signifies descent in general, though in an indeterminate way. For one talks in French of the race of Caesar <in> the same <way> as of the races of horses and dogs, irrespective of the first origin, but, nevertheless, as it seems, always with tacit subordination under the concept of a species... <The word> should mean nothing more than a mass of men whose common formation is distinctive and sufficiently at variance with their neighbors <such that they> could not be immediately derived from them

.

This sounds quite akin to what Hochman would term a "weak concept" and yet this is how race was frequently understood. By this same 'historical consistency criterion' (HCC), we can rule out Hochman's equation of races with taxa subspecies. For one, the race concept preceded the taxa subspecies. For another, nested races were always recognized and yet nested taxa (in the sense of a division and a subdivision of that) can not be assigned to one and the same taxonomic rank. Thus not all races could be assigned to the subspecies category; thus race can not be equated with taxa subspecies unless we wish to disallow nested races.The HCC likewise works against general concepts of race as "large clusters of people that are principally homogeneous within and heterogeneous between" since this is not how races as such were conceptualized either in the 18th, 19th, or 2Oth century. This is not to say that races which represented large clusters of people were not recognized; it is to say that race was used also to describe smaller groups. A quick Ngram search for "microgeographic" and "local" race verifies this point. As shown previous, the HCC rules out all race concepts which require discontinuities and high levels of homogeneity.

Now the HHC might rule against Dobzhansky's broad construction of his Mendelian population concept. But, if so, one can easily salvage the idea or something similar to it by simply only recognizing as races visibly different populations. To set the bounds for which populations should be recognized, one would just look at which were from the mid 1700s to the early 20th century. A quick look at the literature shows that local and even tribal races were frequently discussed, both of which fall fairly far down the scale of differentiation. In short, not much would be lost by restricting the race concept to populations which exhibited differences of magnitudes similar to those that existed between those populations which were frequently called races prior to the 20th century. Likewise the HCC might rule against my natural divisions concept insofar as it claims to be a very general one -- as race was indeed used in an even more expansive and less precise sense e.g., "forms which propagated themselves". But here again there is an easy remedy. Whereas, one can readily constrict Dobzhansky's broad concept to bring it in line with pre-20th century use, one can easily expand mine, just subsuming it under an even more expansive notion of biological race. In my paper I note that this is possible.

My concept works because it corresponds with an actual historical concept which was frequently employed by major players in anthropology and biology and because it happens to more or less correspond with some widely used contemporaneous concepts e.g., "biogeographic ancestry groups", ones which many people recognize as being very race-like. It also works because I adopt pluralism. My concept doesn't preclude others. It either treats them as narrow definitions or as different renderings. On the other hand, the critiques above fail because they cherry pick narrow definitions of race, when not just specific applications of these definitions, and claim that race "really" meant only this, when it is easy, with archives just a few mouse clicks away, to demonstrate otherwise, as I have done across the last 20 to 30 pages of discussion.

Race is a superficial construct. There are too many ethnicities to categorize the entire human population into mongoloid, caucasian, negroid and other flawed racialist terms. There are too many differences between a Dutch and a Polish individual as there are between English and an Italian individuals for them to be all categorized as caucasian.

There are too many ethnicities to categorize the entire human population into mongoloid, caucasian, negroid and other flawed racialist terms. There are too many differences between a Dutch and a Polish individual as there are between English and an Italian individuals for them to be all categorized as caucasian.

Might you clarify the argument? Are you contending that one can not carve out natural (i.e., genealogical-based) human divisions? Or that such would not constitute "races" in the Buffonian or Darwinian sense? Or that such would not correspond with traditional macro-delineations such as "Negroid","Caucasoid" and "Mongoloid"?

Race is a superficial construct.

In what way is the construct qua construct superficial? Do you think that the principle races of Struthio camelus are superficially different?

The article's title has been changed, upon author's request to: "The Nature of Race: the Genealogy of the Concept and the Biological Construct’s Contemporaneous Utility".

This change was due to the paper not showing up on Google Scholar, possibly due to other drafts with the same name being indexed.

Human races are like different colored cats. Despite the difference in visible phenotypes, they are all still cats. Any label to categorize them by color class is superficial.

Human races are like different colored cats. Despite the difference in visible phenotypes, they are all still cats. Any label to categorize them by color class is superficial.

Different colored cats -- of the same species -- could be either morphs (defined in terms of single or complexes of phenotypic differences) or breeds (i.e., domestic races), which are defined in terms of lineage. An example of a complex (di)morphs is male / female. I don't see why you would consider that categorization to be superficial.

The categorization is superficial because there is no use of it. Pets are categorized as breeds because customers sometimes prefer one breed over another.

It is not that people have denied racial differences (they are differences in every individual) but rejected the scientific categorization of racial differences in intelligence capacity simply due to racial differences in socioeconomic and culture factors. It's like comparing apples to oranges.

And applying science to race has no benefit because it's not like gender which will exist forever. Races will eventually intermix and cease to exist as a concept sometime in the distant future.