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Taxonomy as a Rigorous Science

Dinwar

Penultimate Amazing
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Jul 20, 2010
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Someone in this thread made comments about the lack of rigor in paleontology, specifically in regards to taxonomy. I made a brief post in that thread, and I'd like to continue my discussion without derailing the thread.

If I may jump in where I more or less left off:


Scientists have an almost pathological aversion to the subjective, and taxonomists are no different. They have established several methods for ensuring objectivity in their analysis.

First and most importantly is the type specimen. This is a specimen (or more rarely a series of specimens) that defines the species in question. The concept is a bit Platonic/"shadows on a cave wall" for my taste, but even I acknowledge that the function it serves is critical. That function is to give a universal and almost always unverying starting point for species determination. Everyone uses the same type specimens for each species; you can't NOT use the same type specimens. These specimens are carefully maintaned in very high-quality labs. The existence of these type specimens does not, I hasten to emphasize, mean that the concept of a species is immutable once established; any time spent reading the taxonomic literature will show innumerable discussions that amount to "This species should be redefined" (particularly now, with the whole Ceratopsian thing going on). Type specimens make the discussion objective because they give us all a starting point, and all the same starting point.

One of the great tragedies of World War 2 was the bombing of so many museums. I'm not trying to deminish the astounding loss of life; I'm just saying that a tremendous amount of scientific data was irrevocably lost. Unfortunately, this was what was required to spur people into routinely including photographs with their species descriptions. Sadly, many pre-war species no longer have type specimens due to them simply being blown up and converted into rubble. I've had personal experience with the tremendous, and sometimes insurmountable, difficulties this causes even today.

The species description is another way to add objectivity. As I said previously, these descriptions are long, dense, and extremely technical. That's because they need to be. These descriptions give us something against which to test our specimens--they in a very real sense are the formal experiment in taxonomy. Once written, anyone can (once they've been trained to comprehend the description) determine if a new specimen is part of that species or not, using precisely the same logic as physicists used to determine if they'd found the Higgs Boson or not. This means that taxonomy is an empirical, experimental science.

Finally, all of the rules for species identification have been codefied in the International Code of Zoological Nomenclature. This is the framework within which taxonomists are required to operate. The Code defines the different types of type specimens (haplotype, paratype, holotype, and a few more that I've never seen in person), as well as how to address synonymies, contradictions, and other issues that will inevitably arise when thousands of researchers contribute to a conversation that's been going on for three centuries. These rules are determined by the researchers for the purpose of ensuring objectivity.

There are some problems with this method, yes. As I said before, it's very Platonic. The type specimen is supposed to represent the big-T Truth of the species, or at least was until fairly recently. This obviously contradicts evolution, and that assumption was abandoned by most researchers a long time ago. At this point, as far as I can tell, most researchers view type specimens as vital conveniences, not as Horse. I know two researchers (three? depends on how you count them--one person is only intermittently part of that team) who are attempting to redefine ammonite species based on population statistics, rather than a type specimen.

Secondly, and far more significantly, the literature is extremely widely scattered. A common type of taxonomic paper is "A revision of the genus ____", in which the researcher puts together everything they can find about that genus and proposes a new understanding of it. A common criticism of such papers is "But you haven't examined THIS paper!" A valid criticism--and drawing such criticisms is one of the critical aspects of such papers. Unfortunately there is no database we can go to to find this stuff. Several have tried to establish one (I think the Tree of Life is ongoing, but I could be wrong), but the task is beyond the comprehension of most. Even putting together all the information on each family within one order is an enormous task beyond all but the most dedicated research teams. What that means is that, unfortunately, when we describe a new species it's often already been described before. Nothing's more disheartening then realizing that your species was already named, back when scientific publications were in Latin (though it does give you a real sense of your connection with the great names in science; referencing Linnaeus is both disheartening and profoundly satisfying).

Another major issue with taxonomy, and in fact the biggest one, is that it does not presume to define evolutionary history. Linnaeus had no intention of discussing--nor, indeed, any knowledge of--evolution. The book "Darwin's Century" discusses some doubts he had later in life, but when he started at least the nested heirarchy was simply a matter of convenience. This is why Arthropoda can be both polyphyletic and a valid taxonomic name--taxonomy, at its start, was merely about putting things in categories. Which means it has a curious side benefit: taxonomy up until very recently served as a truly independent test for evolution. The fact that taxonomy largely matched evolutionary history, despite having nothing to do with it, is very clear evidence that evolution is right.

Still, taxonomy is a powerful tool. And the thing that strikes me about each of these problems is that the people identifying them and addressing them are researchers in the field. Like Creationists, those who attack paleontology for lacking rigor have not, to my knowledge, presented a single criticism that my coleagues and I haven't addressed first, better, in more detail, and found ways to fix.

-------------------------------

In the other thread I had mentioned that the math behind cladograms using morphological characters and genetic characters (characters=traits; don't ask me why, the best explanation I ever got was "Cladists wanted to set themselves apart, and jargon was an easy way to do that). I'm not going to go into too much depth here; if you really want to know more about the math, here is a very good resource. You can also download a freeware program at that site and actually play with some data yourself. Apparently a new version is out; hopefully they made it so you can copy and paste from Excel into it now. That was one of the most annoying aspects of that program. I'd recommend it to anyone who wants to toy with statistics--it can do all kinds of things, from cluster analysis to PCAs to cladograms to simple Gausian statistics. You just have to play around with it until you find out how. It's for serious researchers; they didn't design it to be intuitive to non-experts.

Anyway, here's how the math works, at the 1:1,000,000 scale: You pick your taxonomic groups, and you pick your characters. Genetic taxonomy is easy--you pick the gene, and each codon or base pair represents a character. Morphological data are trickier, but generally by the time you're doing this stuff you've gained a pretty good understanding of taxonomy so it's not too hard. What you want to focus on are shared, derived traits--meaning traits a group has because their ancestor had it and they retained it. Traits that only individual species have are pretty useless for determining evolutionary history--you already know where they evolved, so they tell you nothing. You should also have an "outgroup", a species that is outside the group you're interested in, but close enough to share some primative traits of the group (bacteria are a horrible out-group for primate cladistics; bats aren't too bad, actually). Then you run the program (it may shut down your computer for a while--on a really big database I saw it shut one down for three days).

What the computer does is construct every possible cladogram and count the number of evolutionary changes necessary to arive at that tree. The program will give you the trees with the elast number of changes necessary, on the assumption that the fewer changes necessary the more likely it is that the tree represents the true evolutionary history (as my old professor put it, "We assume that evolution is hard").

What you will immediately find is that there are many. The reason is that the way the heavy math works is by assuming that only pairs are possible--one species splits into two species, but never three or more. Obviously this doesn't represent biological reality; however, the consensus tree--meaning the average of the shortest trees--typically collapses many of the clades, meaning that it groups multiple species together. There is some debate as to what this means; personally, while I haven't run any serious tests of it (I'm still toying with how to run such a test), I lean towards the idea that those collapsed clades represent evolutionary reality: they show organisms that arose from the same stock, if not necessarily the same time.

Now you can test it. There are multiple ways; the ones I'm most familiar with are bootstrapping and jack-knifing. In one (and I always forget which), you simply remove random characters, re-run the analysis, and see what pops out. It does this a thousand times (default setting; you can make it do more). More robust trees can handle the loss of some data; if, however, some structure relies on a single character, it's more likely that it's not real. Often there's a reporting limit of 50% or so, meaning that if the structure doesn't appear on 50% resultant trees, you treat it as if it's not there. The other test also removes random traits, but it duplicates random traits as well--so each analysis has the same number of characters, but not the same characters. It's dealt with the same way.

This is how the math works for ANY set of characters. Genetic or morphological, even behavioral or stratigraphic (yes, I've seen it; no, I do not approve of if)--the characters don't matter, that's how the math works. So in essence, genetic cladograms are not superior to morphological ones.

One thing that's really cool about these trees is that each time the branches come together (called a node), the math will tell you what the traits of that node are. This forms a hypothesis about the nature of the ancestor of those two species. I've found, through analyzing numerous cladograms, that often the cladogram is saying "This species directly gave rise to this other one", which is by itself a really cool concept (I've toyed around with somehow addressing the distance from nodes, but haven't done much with it since grad school; haven't had time or a reason to get back into it). If the node matches a species you analyzed, that means that you are arguing that that speceis gave rise to the other(s) branching off that node. If the node's characters don't match anything you've analyzed, that means you are hypothesizing the existence of an organism ancestral to the ones you studied, and--here's the part that I find unbelievably awesome--you are predicting the nature of that organism. What that means is that someone can go out, find it, and say "Hey, this matches what they predicted"--making cladistics indespensable for making sense of the past. (The researcher would still have to draft a species description and curate the type specimen; the honor of naming the species goes to the discoverer, not to the person who hypothesized tis existence.)

These mathematical methods have been tested. For example, there was a scientist who created a fictional clade of organisms based on a known evolutionary history. He then presented them to scientists and challanged them to reconstruct the history of that clade. Eventually they succeeded--and we've gotten much better at it over time. As an aside, I want to make some of these as stuffed animals for my impending child.

A second test is to compare the hypothesized evolutionary history against what we see in the rock record. We expect some range extensions--FADs and LADs are not usually the first organism to have evolved in that species, but only the first and last APPEARANCE of that species in the fossil record. Still, the fossil record is useful. If you hypothesize a huge number of organisms and we only find three, or hypothesize traits we just don't find, it's cause to re-examine the cladogram, and specifically the characters and assumptions that went into it.

As I said, this is the 1:1,000,000 scale, for all of this--meaning I'm glossing over a huge amount of data and only barely scratching the surface of the stuff I'm adressing. Still, I think it's sufficient to demonstrate the rigorous nature of taxonomy and of our methods for understanding evolutionary history. If anyone would like me to expand on any of these topics, please feel free. And Jodie, please point to specific areas where subjectivity can be problematic.
 
Very interesting stuff. I'll probably have to read through it a few more times to actually understand it (somewhere after the dotted line my brain started going "bwuh?") :D

It never occurred to me that taxonomy worked as a test for evolution--that's neat!
 
Akri said:
(somewhere after the dotted line my brain started going "bwuh?")
Don't feel bad--my brain pretty much melted each time I attended my Statistical Methods in Paleontology class. The friends I hung out with used to worry about me--I'd spend about an hour staring into space, mumpling things like "But how does an n-dimensional football rotate?" and "But that doesn't prove ancestry!" :D

And my professor was evil. Very wise, but not nice at ALL. She made us work out a cladogram by hand. A small one, and a simple one--but she wanted us to know what was in that black box of a program before we used it. There's a reason cladistics didn't start until a few decades ago: prior to that, humans just didn't live long enough.

If you REALLY want to melt your brain, the journal "Paleobiology" published a paper this year discussing the importance of using random walks as a null hypothesis against which to test paleoecological PCAs. Randomness, multivariat statistics, 26-dimensional space, and paleoecology--this stuff is getting into theoretical physics territory in regards to how much the math hurts your brain, and adds all the issues of taphonomy, collection bias, and ecological reconstruction on top of it! :D
 
Hey Dinwar, what I had in mind is where a species shows a mix of traits. Since Giganto was the original species that sparked the conversation I'll start with that one. As far as I know, no one has identified what the originating species for Giganto was back in Africa. Based on a few jaw bones and numerous teeth, someone came up with the idea that Giganto was bipedal.

I understand that the assumption rose from how they think the spine attached to the skull, that this was an aspect of bipedalism. So if you used the technology and math you listed above, then I'm assuming they compared that with other bipedal and semi bipedal species to come to that conclusion. I still think it's subjective to make that assumption based on so few examples of bipedal species, namely us and our ancestors. The math and processes are sound but the pool you are pulling the data from is small. That was my point.

The mechanisms underlying population level variation have been shown to be directly related to diversity within the species. In that case that would be useful for evolutionary predictions, but if you don't have that diversity represented in the fossil record because of the limited amount of fossils compared to the overall population through time, how can it be of any use? Are you back engineering your predictions based on modern day genotypes and how they correlate to modern day morphology?
 
I still think it's subjective to make that assumption based on so few examples of bipedal species, namely us and our ancestors. The math and processes are sound but the pool you are pulling the data from is small. That was my point.
I asked you this in the other thread, but it's more relavant here: what sample size is necessary for conclusions to not be subjective or assumptions, and how did you arrive at that number?
 
I do statistics for epidemiology, for example, if I only have 8 Thai people living in a community here in South Carolina and three have TB, is it fair to say that TB seems to be a more prevalent disease found among Thai people as a general rule? It might be true if you go to Thailand and compare rates of incidence with the United States but in paleontology you don't have that kind of pool to pull from.

If you only have six examples of a specific hominid species, how do you derive how large the population might have been at it's peek or figure out what is a normal variation within that group? I understood what Dinwar posted, and was interested in how the predictions were made, but when you start with such small numbers how can you predict anything from that? The only way I can see how you might do it is by comparing it to other hominid species and what is evident in modern man. It still seems kind of subjective to me since you don't have a large group of examples from the ancient species in question for comparison.
 
Anthropologists Confirm Link Between Cranial Anatomy and Two-Legged Walking
http://www.sciencedaily.com/releases/2013/09/130926111903.htm
As part of the study, the researchers measured the position of the foramen magnum in 71 species from three mammalian groups: marsupials, rodents and primates. By comparing foramen magnum position broadly across mammals, the researchers were able to rule out other potential explanations for a forward-shifted foramen magnum, such as differences in brain size.

According to the findings, a foramen magnum positioned toward the base of the skull is found not only in humans, but in other habitually bipedal mammals as well. Kangaroos, kangaroo rats and jerboas all have a more forward-shifted foramen magnum compared with their quadrupedal (four-legged walking) close relatives.
Seems relevant.
 
It is, so they look at what is to determine if the majority have it and decide that it is diagnostic, much like we do with disease. Well how did they do it with just Giganto's jaw?
 
I do statistics for epidemiology, for example, if I only have 8 Thai people living in a community here in South Carolina and three have TB, is it fair to say that TB seems to be a more prevalent disease found among Thai people as a general rule? It might be true if you go to Thailand and compare rates of incidence with the United States but in paleontology you don't have that kind of pool to pull from.

If you only have six examples of a specific hominid species, how do you derive how large the population might have been at it's peek or figure out what is a normal variation within that group? I understood what Dinwar posted, and was interested in how the predictions were made, but when you start with such small numbers how can you predict anything from that? The only way I can see how you might do it is by comparing it to other hominid species and what is evident in modern man. It still seems kind of subjective to me since you don't have a large group of examples from the ancient species in question for comparison.
This doesn't answer my question. In order to say "this sample size is too small" you have to know what an appropriate sample size would be. Obviously bigger samples are always better, but you're arguing that there's a lower limit at which point the sample size is ineffective. I'm asking you to say exactly what that limit is (if you can't then you can't say a sample size is too small) and how that specific number was derived.
 
Anything involving any amount of guesswork is pretty much always going to be at least a little subjective. I mean, the facts the guess is based on are objective, but the implications are going to be open to some amount of interpretation, which is why scientists generally use phrases like "this suggests" and "this finding may indicate" or, when there's "a lot" (also a subjective judgment) of evidence "it is likely".

This is kind of related to, but somewhat different from the "species problem" as I understand it, though.
 
This doesn't answer my question. In order to say "this sample size is too small" you have to know what an appropriate sample size would be. Obviously bigger samples are always better, but you're arguing that there's a lower limit at which point the sample size is ineffective. I'm asking you to say exactly what that limit is (if you can't then you can't say a sample size is too small) and how that specific number was derived.

All I can look at is the estimated population of the species that used to exist versus what we actually have in the fossil record. You could use Neanderthal as an example.

I think they estimate the Neanderthal population at 70,000 at its peak. I think bones from about 3-400 have been found, I'm not sure if they have any complete skeletons. That doesn't seem like enough to me to draw any kind of conclusion if I were trying to map health statistics without knowing what was going on with the other 65,000 members of the population. However, unlike many of the ancient hominids, at least in this case they do have DNA.
 
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All I can look at is the estimated population of the species that used to exist versus what we actually have in the fossil record. You could use Neanderthal as an example.

I think they estimate the Neanderthal population at 70,000 at its peak. I think bones from about 3-400 have been found, I'm not sure if they have any complete skeletons. That doesn't seem like enough to me to draw any kind of conclusion if I were trying to map health statistics without knowing what was going on with the other 65,000 members of the population. However, unlike many of the ancient hominids, at least in this case they do have DNA.
I've bolded the most problematic part of your post. You don't have any kind of objective answer to the question "how small of a sample size is too small, and why?" You just have what "doesn't seem like enough" to you.

In other words, your complaint that paleontology is subjective is based on your own subjective idea of how big a sample size should be. Pot, meet kettle.
 
Statistics tells you the significance of the sample size, not me. The numbers suggest a sample size of 1800 individuals for that population to have a 95% confidence level for whatever you want to say about them, doesn't mean they aren't right about the Neanderthals, just that you can't make definitive conclusions based on looking at the fossils alone. I'm sure the DNA evidence adds to that.

I guess if you use a survey of paleontologist's opinions on how much variation might exist in that population it might make a difference, or you could factor in how much variation exists within modern homo sapiens using the Neanderthal DNA as a bench mark. It depends on how you want to look at it, which can be subjective.
 
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Statistics tells you the significance of the sample size, not me.
Except that paleontologists don't agree with your assessment about sample size (they don't agree that sample size matter at all, because the type specimen renders sample size moot). So you need to demonstrate why a certain minimum sample size is needed, and how you got that number. And you actually need to demonstrate that not doing things this way is problematic for paleontology. Simply saying that you think it's problematic isn't enough--you need to actually show actual errors being caused by paleontologists not using your statistical method.
 
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When the evidence is lacking, they make guesses and qualify them with statements like this:

http://www.pnas.org/content/87/20/8120.full.pdf
Gigantopithecus has always been an enigma for
paleoanthropologists because its body size and weight almost
certainly far exceed the range of any other primate, living or
extinct (16-18), and it is the only known species of ape to
have become extinct during the Pleistocene.

And going back to the 60's:

http://onlinelibrary.wiley.com/doi/10.1525/aa.1965.67.5.02a00160/pdf

If we accept Pei’s interpretation of the ecology of Gigantopithecus as a hunter of
large ungulates who apparently dragged his prey back to the cave, then this seems to imply that Gigantopithecus was bipedal much of the time. No bones have been found
which would shed light on this question.
 
Except that paleontologists don't agree with your assessment about sample size (they don't agree that sample size matter at all, because the type specimen renders sample size moot). So you need to demonstrate why a certain minimum sample size is needed, and how you got that number. And you actually need to demonstrate that not doing things this way is problematic for paleontology. Simply saying that you think it's problematic isn't enough--you need to actually show actual errors being caused by paleontologists not using your statistical method.

Statistics is a form of math, a tool used by science. If you question the validity of the tool then that affects the validity of all other conclusions, making them subjective.

You take 70,000 as your population total, and get your percentage of that population using 400, which is 5.7% of the population. That percentage gives you an interval of 2.27% and you use that to get the sample size needed from 70,000 to give you a 95% confidence level. In this case it would be 1800. Now if you are doing a survey of attitudes towards a health concern, the population size wouldn't be so important.

I'm not sure how you would apply that to this situation unless you were looking at paleontologist's opinions in which case they all decide that one specimen is all that is needed........so taking one specimen that is all that is needed you determine whether it is/was a different species from modern humans, that doesn't seem like a giant leap to you using your example? I'm just asking.
 
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All I can look at is the estimated population of the species that used to exist versus what we actually have in the fossil record. You could use Neanderthal as an example.

I think they estimate the Neanderthal population at 70,000 at its peak. I think bones from about 3-400 have been found, I'm not sure if they have any complete skeletons. That doesn't seem like enough to me to draw any kind of conclusion if I were trying to map health statistics without knowing what was going on with the other 65,000 members of the population.

I'm fairly sure they haven't even been attempting to precisely map the health statistics?

They say stuff like this:
http://bryanhockett.com/uploads/JQI_2883_Corrected_Proof_Hockett.pdf

The fate of the Neanderthals is known. Migrations of modern
humans into northern Europe commenced about 40,000 years ago.
Approximately 10,000 years later, modern humans had spread
throughout Europe, and the 250,000 year occupation by Neanderthals came to an end.
How the Neanderthals met this fate is a matter of debate.Differences in dietary intake and energy requirements between
Neanderthals and modern humans are possible contributing factors.
 
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Oh I know Kelly, I was just applying the type of statistics I use to the estimated peak population of Neanderthals versus how many fossils of individuals we actually have as the example. The original debate stemmed from my saying I thought paleontology was subjective. Dinwar says it's more rigorous because they have tested statistical means to predict and correlate who belongs in what species.

I'm sure morphology doesn't equate to disease incidence or survey polls. In the case for Neanderthal we do have DNA to back up the morphology in the 400. That 400 are not all identical, some exhibit features that are more modern the closer you approach modern times.

I see what Dinwar is saying about the complexity. My point is that there are general assumptions made with less than adequate samples such as those ancient hominids that we don't have DNA for, but from what I'm gathering they look at morphology across species. I don't see anything mentioned about the possibility of it being co-evolution. You wouldn't necessarily know that the same combination of genes would produce the same features in those different species for which we lack DNA.

Example: A different group of genes is responsible for red hair in Neanderthals than those responsible for red hair in modern humans.
 
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Statistics is a form of math, a tool used by science.
In certain places, where applicable. I'm asking you to demonstrate that this is one of those places. Simply throwing out numbers doesn't cut it--you need to show that the numbers are relevant to paleontology, and that not using them causes problems. If not using statistics doesn't cause any problems (and "it seems wrong to Jodie" isn't a problem) then there's no issue.

I'm not sure how you would apply that to this situation unless you were looking at paleontologist's opinions in which case they all decide that one specimen is all that is needed........so taking one specimen that is all that is needed you determine whether it is/was a different species from modern humans, that doesn't seem like a giant leap to you using your example? I'm just asking.
If I showed you pictures of chimpanzees, what sample size would you need to determine that they were not pictures of humans?
 
My point is that there are general assumptions made with less than adequate samples such as those ancient hominids that we don't have DNA for, but from what I'm gathering they look at morphology across species. I don't see anything mentioned about the possibility of it being co-evolution. You wouldn't necessarily know that the same combination of genes would produce the same features in those different species for which we lack DNA.

Example: A different group of genes is responsible for red hair in Neanderthals than those responsible for red hair in modern humans.

Can you give a specific example of "general assumptions made with less than adequate samples"?

With genetics and Neandertals and hair/skin genes, that's more of a question for geneticists than paleos, but either way, the geneticists say things like:


http://media.longnow.org/files/2/REVIVE/NeanderthalMC1R_Science2007.pdf
The impaired activity of this variant
suggests that Neanderthals varied in pigmentation levels, potentially on the scale observed in
modern humans. Our data suggest that inactive MC1R variants evolved independently in both
modern humans and Neanderthals."

And this:
http://scholar.google.com/scholar?cluster=4360132809304065031&hl=en&as_sdt=0,43&as_ylo=2009

Predicting Homo Pigmentation Phenotype Through Genomic
Data: From Neanderthal to James Watson

The archaic hominid and Paleo-Eskimo phenotypes cannot be validated because they are extinct, so that only speculations can be made (subjects 12–15, 30; Tables 1
and 2). Lalueza-Fox et al. (2007) found a MC1R gene variant (R307G) in two Neanderthals from Italy and Spain,
which could determine pale skin and/or red hair as
observed in modern humans.
 

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