Denisovans and the species problem

Posted 21 January 2011 by

A few weeks ago I blogged on the Denisovans, a new group of human relatives discovered through genetic analysis of two bones from Denisova in Siberia (Reich et al. 2010, Nature 468:1053). Fascinatingly, the Denisovans seem to have made about a 5% contribution to the genome of living Melanesians. I mentioned that this new discovery did not seem compatible with a young-earth creationist framework, and awaited with interest a creationist explanation of the findings. Answers In Genesis (AIG) has now commented on the Denisovans. No explanation, merely a one-sentence handwaving solution:

But the most interesting twist (from the evolutionary perspective) is that modern humans from New Guinea have Denisovan DNA. While an evolutionary perspective interprets this as meaning that Guineans' ancestors "interbred" with Denisovans, a biblical perspective interprets this as simply meaning that the descendants of one of the people groups leaving Babel eventually settled in what is now New Guinea.

— Answers in Genesis
It's not clear what this even means. After all, their 'biblical perspective' had exactly the same interpretation (that the descendants of a group leaving Babel settled in New Guinea) even before we knew about the Denisovan genetic contribution. This 'explanation' fails to address a key point: how did the Denisovan genes get into Melanesians, if not by interbreeding with Denisovans? And the above scenario doesn't resolve any of the other problems with a young-earth framework. Why is the Denisovan genome so different from all modern humans, if they were descended from the same eight people on the Ark? (The Denisovans would, presumably, have to be another group of people who left Babel, since Babel happens after the global flood in the Bible.) Why does Africa have the greatest genetic diversity in modern humans but no Neanderthal or Denisovan genes? Why are Neanderthal genes found in all non-Africans, but not in Africans? The genetic diversity of modern humans, Neanderthals and Denisovans is much greater than that of modern humans alone. How could there be so much genetic diversity so soon after disembArking from the Ark? How did all that extra genetic diversity disappear? Genes can disappear quickly in population bottlenecks, or slowly through random processes like genetic drift in stable populations, but it's very unlikely for genes, let alone large numbers of genes, to disappear in a rapidly expanding population (and going from 8 to 7 billion in under 10000 years is definitely rapid expansion). On to the question of what species the Denisovans should be assigned to. Cautiously, and commendably IMO, the scientists declined to classify the Denisovans taxonomically, given that we know almost nothing about their anatomy. Answers in Genesis, of course, considers them all humans:

Writing for BBC News, Clive Finlayson, director of the Gibraltar Museum, emphasizes that both Denisovans and Neanderthals belonged to our species, Homo sapiens. (Indeed, given the original definition of species as referring to organisms that could interbreed successfully, treating them as separate species doesn't make sense. However, that definition is no longer observed.)

— Answers in Genesis
There's a reason why scientists have struggled since before Darwin to define what a species is. It's called the 'species problem', and it's why so many species definitions and concepts have been proposed. Basically, the world is a complex, messy place. Scientists would love it if they could use AIG's definition. They can't and don't, because it doesn't work in the real world. It may be simple, but it's also simple-minded. Do we really want to put lions and tigers in the same species? Dogs and wolves? The fact that lions and tigers can interbreed is less important to biologists than the fact they they differ in many other important ways. There are other problems. For example, AIG's definition doesn't handle cases where species A can interbreed with B, and B with C, but A can't interbreed with C. It's an inherently difficult problem that will probably never be fully solved because the complexity of life defies easy categorization.

225 Comments

Ron Bear · 21 January 2011

Could someone please explain to me what "the Denisovans seem to have made about a 5% contribution to the genome of living Melanesians" means. I know I am doing something stupid, but I don't know what. We're 98 percent the same genome as Chimps, but Melanesians got 5 percent from Denisovans. Is that 5 percent of the 2 percent that makes us different from Chimps? It is obvious to me that the 98 percent and the 5 percent can't be a percentage of the same thing, but I'd like to understand what I am missing here. Thanks in advance.
Ron

Chris Caprette · 21 January 2011

Answers in Genesis Wrote: (Indeed, given the original definition of species as referring to organisms that could interbreed successfully, treating them as separate species doesn’t make sense. However, that definition is no longer observed.)
This statement from AIG is false (big surprise) on two fronts. First, typological species concepts long predated interbreeding criteria. Second, many biologists still use interbreeding criteria to distinguish species for many, if not most vertebrates. Leave it to the creationists to keep shoveling the manure.

DS · 21 January 2011

Of course the reason for the "species problem" is that descent with modification is true. That is where species come from. That is why they are not nice neat packages the way that creationists want them to be. This is completely consistent with evolutionary theory.

It might be more appropriate to define species by genetic discontinuity, since genetic divergence is an inevitable consequence of reproductive isolation. This approach would also allow for the determination of levels of introgressive hybridization, which seems to be the case here.

Apparently, there was some interbreeding in the past of some Denisovans and some modern humans. Apparently some of these genes, which can be distinguished from the lineage they introgressed into due to the degree of genetic divergence, can still be found in some modern populations. That is where the 5% figure comes from. It doesn't have anything to do with the divergence of humans from chimps, since they last shared a common ancestor before the split of modern humans and Denisovans.

Flint · 21 January 2011

It is obvious to me that the 98 percent and the 5 percent can’t be a percentage of the same thing, but I’d like to understand what I am missing here. Thanks in advance. Ron

Well, see, both the Denisovans and the Melanesians were slightly modified chimps (as are we). But they were slightly modified in different ways. Some of those differences got passed through interbreeding.

Joe Felsenstein · 21 January 2011

Ron Bear said: Could someone please explain to me what "the Denisovans seem to have made about a 5% contribution to the genome of living Melanesians" means. I know I am doing something stupid, but I don't know what. We're 98 percent the same genome as Chimps, but Melanesians got 5 percent from Denisovans. Is that 5 percent of the 2 percent that makes us different from Chimps? It is obvious to me that the 98 percent and the 5 percent can't be a percentage of the same thing, but I'd like to understand what I am missing here. Thanks in advance. Ron
They aren't percentages of the same thing. Human genes differ from each other by about 0.1% (genes in Denisovans presumably differ a bit more from those in modern humans). The 5% figure is roughly the fraction of genes (loci) at which the small differences between different copies indicate that Melanesian copies are more similar to Denisovan copies than they are to copies in other humans.

RodWl · 21 January 2011

We teach the various definitions of a species in our 100-level Bio course. I always thought that the viable-offspring definition was the consensus view since its the least arbitrary but then Neanderthals wouldnt be a separate species.
I suspect that any other 2 organisms separated by the equivalent time/genetic distance/morphology would be considered the same species and its just our human bias that splits them into another species

Ron Bear · 21 January 2011

Thanks for explaining. So Denisovans contributed roughly .05 percent of the entire genome of Melanesians. That makes it so that the math squares up on my original question. Also it makes sense to me that if someone is trying to determine how I am related to you or how either of us is related to a Denisovan, it wouldn't make sense to look at the entire genomes. It makes sense to look at the differences between the genomes.
Thanks,
Ron

John Harshman · 21 January 2011

RodWl said: I always thought that the viable-offspring definition was the consensus view since its the least arbitrary but then Neanderthals wouldnt be a separate species.
That's the trouble with species definitions. Not a one of them doesn't encounter problems here and there. The viable-offspring definition is as arbitrary as any. How, for example, does it deal with cases in which male-A female-B crosses are viable but female-A male-B crosses are not? Or cases of simply reduced numbers of viabile offspring? Or, closer to my heart, how would it deal with the family Anatidae, which under the viability definition would be approximately a single species, there being known crosses between subfamilies and between most genera?
I suspect that any other 2 organisms separated by the equivalent time/genetic distance/morphology would be considered the same species and its just our human bias that splits them into another species
Sorry, but no. Simple genetic distance doesn't make a species. Nor does morphology. We might make a guess on either basis, but the guess doesn't have to correspond to reality. If there's a consensus view (and there probably isn't), the question to ask is whether the two populations would coalesce into one population if they were sympatric. Easy to answer for populations that actually are sympatric, harder for those that are allopatric. Consider the example of the rosy finches (Leucosticte). They differ enough in plumage that it's easy to tell them apart. They don't differ at all genetically, so far as can be told, in that mtDNA haplotypes are randomly shared among populations. (Obviously, they must differ, just not at loci we've examined so far.) And their breeding ranges don't overlap, though their winter ranges do. Anyway, the point is that they are genetically much less differentiated than Denisovans and modern humans, but we do call them good species.

Jeremiah Tattersall · 21 January 2011

I've talked with my local open air creationist preacher about the new finding about Denisovan (we have weekly coffee then screaming matches). After we got through all the BS about dating methods and sequencing his argument position was that of Magic Man done it.
To quote: "If the fossils seem old it's because god made them seem that way. If the DNA seems like it was sequenced it's because god made it seem that way, not because it actually is that way".
That's right. God, the great deceiver.

Jim Thomerson · 21 January 2011

Your open air creationist is indulging in what has been called the "deceptive God blasphemy".

There is a good bit of species criteria discussion on the "Why Evolution is True" blog, centered on how many elephant species. I've studied interspecies hybridization some, and have just been reading a manuscript about why two sister species hybridize here, but not there, and what it all means. I suspect the large majority of sexually reproducing biparental animal species are well isolated genetically out in nature. Instances of interspecies hybridization are, I think, relatively rare and therefore interesting as a situation which tests the theory of species being genetically isolated.

raven · 21 January 2011

Cautiously, and commendably IMO, the scientists declined to classify the Denisovans taxonomically, given that we know almost nothing about their anatomy.
That is what I'd really like to see. What do the Denisovans look like? A few skulls would be nice. Presumably they would look vaguely like sapiens sapiens and sapiens neaderthal but the details are anyone's guess right now. IIRC, one of the artifacts they found in the same cave was a bracelet. A group that can survive in Siberia and make jewelry doesn't sound too dumb.

Marion Delgado · 21 January 2011

This is basic Creation Science! There were giants in the earth in those days; and also after that, when the sons of God came in to the daughters of men, and they bore children to them, the same became mighty men which were of old, men of renown.

Marion Delgado · 21 January 2011

Twelve thousand years old. But I actually asked this guy, "OK, dinosaur fossils-- how does that fit into your scheme of life? What's the deal?" He goes: --"God put those here to test our faith."
--"I think God put you here to test my faith, dude. I think I've figured this out."
Does that-- That's what this guy said. Does that bother anyone here? The idea that God might be @#$%ing with our heads? Anyone have trouble sleeping restfully with that thought in their head? God's running around burying fossils: "Ho ho! We'll see who believes in me now, ha ha! I'm a prankster God. I am killing me, ho ho ho!"
You know? You die, you go to St. Peter:
"Did you believe in dinosaurs?"
"Well, yeah. There were fossils everywhere. (trapdoor opens)
Aaaaarhhh!"
"You #$@@$#in' idiot! Flying lizards? You're a moron. God was %$#%#in' with you!"
"It seemed so plausible, aaaaaahh!"
"Enjoy the lake of fire, @#$@er!"
--- Bill Hicks

John Harshman · 21 January 2011

You don't see the "test our faith" trope very often these days, probably because of the obvious theological problems. Then again, the story of Abraham and Isaac would seem to argue in favor of such a god. Not to mention that in the Eden story, the snake was telling the truth and god was lying about the tree.

raven · 21 January 2011

JF: Human genes differ from each other by about 0.1%
http://www.sciencedaily.com/releases/2007/09/070904072204.htm. (from 2007) While the SNP events outnumbered the non-SNP variants, the latter class involved a larger portion (74%) of the variable component of Dr. Venter's genome. This data suggests that human-to-human variation is much greater than the 0.1% difference found in earlier genome sequencing projects. The new estimate based on this data is that genomes between individuals have at least 0.5% total genetic variation (or are 99.5% similar) The researchers suggest that much more research needs to be done on these non-SNP variants to better understand their role in individual genomics.
genome.gov: Genetic Variation Program Overview Most of any one person's DNA, about 99.5 percent, is exactly the same as any unrelated person's DNA. Differences in the sequence of DNA among individuals are called genetic variation.
FYI. The 0.1% difference is from older estimates. From whole genome sequencing, it is now given as 0.5%, 15 million bp different. Which isn't all that much less than human chimpanzee given as 98 or 99%. This has always implied (IMO) that which sequences are different and why is more important than the absolute numbers.

John Harshman · 21 January 2011

Raven, not that I doubt you, but can you give a citation for that estimate, and does it use the same index of similarity as the human-chimp figure of 1.3%, based on site differences in aligned sequences?

raven · 21 January 2011

Raven, not that I doubt you, but can you give a citation for that estimate, and does it use the same index of similarity as the human-chimp figure of 1.3%, based on site differences in aligned sequences?
Ummm, I did. There is a link in that post to the sciencedaily.com article from 2007 at the top of the box. Typing a few keywords in google including "99.5%" will bring up the info from genome.gov. AFAIK, the human human and human chimpanzee differences use the same type of sequence analysis.

OgreMkV · 21 January 2011

I'm not a practicing Biologist, so I can safely say that I don't think species exist. Within a genera (or possibly a larger group, thanks John), it's just a continum.

Humans like to be able to put things into neat boxes, but clines just don't allow organisms to fit into neat little boxes. They have to be really big boxes, and then what's the point of putting them into boxes.

psweet · 21 January 2011

Jim Thomerson said: Your open air creationist is indulging in what has been called the "deceptive God blasphemy". There is a good bit of species criteria discussion on the "Why Evolution is True" blog, centered on how many elephant species. I've studied interspecies hybridization some, and have just been reading a manuscript about why two sister species hybridize here, but not there, and what it all means. I suspect the large majority of sexually reproducing biparental animal species are well isolated genetically out in nature. Instances of interspecies hybridization are, I think, relatively rare and therefore interesting as a situation which tests the theory of species being genetically isolated.
You may well be right about the 'large majority' of species being well isolated (most of our animal species are arthropods, and I don't have much of a feel for them). But there are a few groups that really make the problem stand out. John Harshman mentioned the Anatidae, I can only guess he's scared to mention the Laridae. Trying to sort out the taxonomy of the LWHG (Large, white-headed gulls) seems to be a true nightmare. Hybridization is actually very common in places and in some populations, to the extent that the majority of birds in some locales are considered to be interspecies hybrids. (The entire coast of Washington state, for example.) Yet we can clearly diagnose many of these same species when they wander to other locations. (Well, maybe not as clearly as we'd like to think.) I think that the Anatidae includes a couple of cases that indicate how much historical biases can get in the way of our classification, specifically the Hawaiian Duck and the American Black Duck. Both have historically been considered full species, and both are in the process of being 'genetically swamped' by Mallards. I'm not sure if there is a valid argument for maintaining their species status, other than conservation concerns.

david winter · 21 January 2011

Excuse me for the shameless self-promotion. But I think the evidence that modern humans population derrive genes from Neanderthal and Denisnovans is a really good way to edge into the species problem.

As others have said, the difficulty we find in drawing a bright white line between species is actually a prediction of descent with modification and something of a problem for creationists who want to separate our species from the rest of creation.

Mike Elzinga · 21 January 2011

david winter said: As others have said, the difficulty we find in drawing a bright white line between species is actually a prediction of descent with modification and something of a problem for creationists who want to separate our species from the rest of creation.
I guess ID/creationists just have to accept the fact that bread dough can’t turn into bread and cake batter can’t turn into a cake.

Jim Thomerson · 21 January 2011

OgreMK5, I'm a retired fish alpha taxonomist. I spent my career identifying fish species and considering their relationships at the generic and family levels. This includes describing new species and genera, and producing keys for species identification. My experience does not support your idea that species ordinarily form continua.

mrg · 21 January 2011

Mike Elzinga said: I guess ID/creationists just have to accept the fact that bread dough can’t turn into bread and cake batter can’t turn into a cake.
Oh, but those are the product of an Intelligent Baker.

Alan Barnard · 21 January 2011

I am not a biologist so I can give you no insight into what a species is. But why do you want to introduce a term for a quality that you cannot define? Compare it with the concept of aether, we did away with this, once we understood what was actually happening, without any pain.

We have known what was actually happening in biology for even longer. Darwin drew his 'I think' tree and we have never looked back.

If we could zoom in on the tree so as to see individual creatures, we would find that each twig consisted of myriads of little lives which intersect and branch while remaining part of the twig. Sometimes the twig will bifurcate and the two parts go their separate ways. Highly magnified, it may not be possible to tell exactly where the separation occurred or where it became irrevocable (where no lives are able to cross from one to another or, if they do, fail to produce fertile offspring).

Knowing what we do, is it sensible to try to cut some slice out of a branch of the tree and say that this is a particular 'species'?

Mike Elzinga · 21 January 2011

mrg said:
Mike Elzinga said: I guess ID/creationists just have to accept the fact that bread dough can’t turn into bread and cake batter can’t turn into a cake.
Oh, but those are the product of an Intelligent Baker.
Not when I do it. :-)

mrg · 21 January 2011

Alan Barnard said: Knowing what we do, is it sensible to try to cut some slice out of a branch of the tree and say that this is a particular 'species'?
Yes. Are cats and dogs different species? Unambiguously. How about housecats and caracals? That's more ambiguous. Are German and French different languages? Unambiguously. Are French and Walloons different languages? That's more ambiguous. We define different colors, but anyone looking at a color wheel might have some difficulty determining where one begins and another leaves off. The concept of a species is not so different in concept than the concept of a color, and I don't see that it would be useful to abandon either concept simply because they are ambiguous. The real world is like that.

mrg · 21 January 2011

Mike Elzinga said:
mrg said:
Mike Elzinga said: I guess ID/creationists just have to accept the fact that bread dough can’t turn into bread and cake batter can’t turn into a cake.
Oh, but those are the product of an Intelligent Baker.
Not when I do it. :-)
I can sympathize. When we get to a certain age, it's difficult to make things rise any more.

Mike Elzinga · 21 January 2011

mrg said:
Mike Elzinga said:
mrg said:
Mike Elzinga said: I guess ID/creationists just have to accept the fact that bread dough can’t turn into bread and cake batter can’t turn into a cake.
Oh, but those are the product of an Intelligent Baker.
Not when I do it. :-)
I can sympathize. When we get to a certain age, it's difficult to make things rise any more.
Ah; Viagara instead of yeast in the bread dough! I hadn’t thought of that.

mrg · 21 January 2011

Mike Elzinga said: Ah; Viagara instead of yeast in the bread dough! I hadn’t thought of that.
I was thinking of several rejoinders to that, but they tend to accelerate the descent of the conversation rapidly.

OgreMkV · 21 January 2011

Jim Thomerson said: OgreMK5, I'm a retired fish alpha taxonomist. I spent my career identifying fish species and considering their relationships at the generic and family levels. This includes describing new species and genera, and producing keys for species identification. My experience does not support your idea that species ordinarily form continua.
Just out of curiosity, did that include cichlids?

Mike Elzinga · 21 January 2011

mrg said:
Mike Elzinga said: Ah; Viagara instead of yeast in the bread dough! I hadn’t thought of that.
I was thinking of several rejoinders to that, but they tend to accelerate the descent of the conversation rapidly.
Yeah; to say nothing of the irony of having it all ending up on the Bathroom Wall.

Jim Thomerson · 21 January 2011

http://books.google.com/books?id=sltlCl1XgJwC&printsec=frontcover&dq=Belize+Thomerson&source=bl&ots=3KDj5NqlCN&sig=r6gcD3Eg6WWZuxUTMQNqylaF0Fk&hl=en&ei=ZBc6TeqEJcO78gakufiPCg&sa=X&oi=book_result&ct=result&resnum=1&ved=0CBMQ6AEwAA#v=onepage&q&f=false

We identified 15 species of cichlids from Belize. We identified Petenia splendia, and left the other 14 species in Cichlasoma, knowing that revisions of the new world cichlids were underway, and probably most or all of them are no longer in Cichlasoma. C. maculacauda occurs in southern Belize, and the very similar C. synspilum in northern Belize. I suspect additional collections in central Belize might uncover a cline, but we did not have specimens to support anything more than vague suspicions.

mrg · 21 January 2011

Mike Elzinga said: Yeah; to say nothing of the irony of having it all ending up on the Bathroom Wall.
Oh, that's free-fall all the way to the basement.

John Harshman · 21 January 2011

OgreMkV said: I'm not a practicing Biologist, so I can safely say that I don't think species exist. Within a genera (or possibly a larger group, thanks John), it's just a continum. Humans like to be able to put things into neat boxes, but clines just don't allow organisms to fit into neat little boxes. They have to be really big boxes, and then what's the point of putting them into boxes.
"Exist" is such an problematic word. But nevertheless, I think species do exist. No, genera aren't continua. Diversity is clumped, and there are few smooth clines between populations. (There are some.) There is structure within species, and there are various degrees of interbreeding between things we call species, and both of these muddy the matter a bit. But by and large, the concept of species reflects the way nature works pretty well. At least for obligately outcrossing eukaryotes.

John Harshman · 21 January 2011

Alan Barnard said: Knowing what we do, is it sensible to try to cut some slice out of a branch of the tree and say that this is a particular 'species'?
Well, most of what we're looking at is the very tips of the branches. The concept of species actually works pretty well if you restrict it to organisms in one time-slice (usually the present). It can start to have problems if you try to extend it over evolutionary time. Similarly, by the way, it works really well over a limited geographic area, and can (though less often) start to have problems when you extend it over continental ranges. But in general, we use species because they encapsulate a very important feature of the non-uniform distribution of diversity. If you take the color analogy, we don't in fact see anything analogous to a continuous spectrum. We see a range of frequencies we might call "red" that's separated by a gap from some range of frequencies we call "orange", and so on. There are of course cases where, say, blue shades into green. But not so many as to make the concept useless.

mrg · 21 January 2011

John Harshman said: No, genera aren't continua. Diversity is clumped, and there are few smooth clines between populations. (There are some.)
I remember going through Darwin's ORIGIN OF THE SPECIES and running into a discussion in chapter six that started off:
Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion instead of the species being, as we see them, well defined?
Well ... relatively well defined. Anyway, it took me a while to dope out what he was talking about: http://www.vectorsite.net/taorgin_03.html#m1 Speciation could be thought of as differing historical trajectories of two related populations that have become isolated from each other.

JGB · 21 January 2011

As luck would have it had introduced the species problem and such to my biology class in early December starting with a collection of human skull images that have been typed by race. I ask the students to naively classify what they see into species groups. After they come up with their ideas and debate a bit I reveal my chicanery, and then present them with a randomized image of the hominid skulls from the Smithsonian. It forces them to confront the phenotypic similarity in a rather visceral way. Fast forward a few weeks and they had read and discussed the Chris Stringer lecture from the Talk Origins archive, and discussed the relative humanity of Neanderthals right before winter break. I was then able to discuss after break the serious implications for the Neanderthal and Denisovan DNA sequencing. I felt that those sequences were much better at reinforcing the application of the species problem to humans, rather than as an introduction, which was suggested above.

John Harshman · 21 January 2011

raven said: AFAIK, the human human and human chimpanzee differences use the same type of sequence analysis.
It turns out they don't. The intra-human comparison takes into account indels, including copy number differences. By that measure, humans and chimps are about 95% similar. I think this is a silly way to count genetic distance, by the way. But anyway, it's either 99.9% vs. 98.7% or 99.5% vs. 95%. Take your pick, but don't mix and match.

Jim Thomerson · 21 January 2011

20th century biologist me has been thinking about the Phylogenetic Species Concept. Is this a reasonable statement of the concept? A species originates at a speciation event and exists through time until it becomes extinct. This adds a time, historical, dimension to the the species which we don't see when we look at species, as we do, in a little slice of time. So what we see, in the moment, is a Biological Concept Species or whatever. I have the feeling I am missing something here.

Henry J · 21 January 2011

Identifying the exact point at which a species branches off from another is a bit like identifying the precise boundary between a tree branch and the tree trunk (or its parent branch).

Henry

Leszek · 22 January 2011

I am not sure this helps, but I heard this once on Canadian TV and it sticks with me because it helped drive it home. I also wonder what you guys here think of this:

Lets imagine we are standing face to face with a modern chimpanzee. We hold that hand of our mother and her other hand is holding the hand of her mother. So on it goes down the generations. Opposite us the chimp is doing the same. With an average separation of 1 meter (just over a yard for you non metric types) a line would stretch from Windsor (Canadian side of the boarder from Detroit) down the line until at around Toronto something interesting would happen. The Chimp line and your line would be holding the opposite hands of the same mother.

If you stretch these two lines out to one continuous line, there is no point at which you can say everyone on this side is Human and everyone on that side is Chimp without breaking up a mother and child. It is only easy to identify one or the other because all the ancestors are already dead. The concept of species works really well in those situations.

But for some species such as ring species not all the intermediates are dead. Some of them have dead intermediates but the species themselves are still closely related enough that they could interbreed, even if poorly. There are all sorts of variations in between.

I hope that makes it clear or at the vary least is a new way to look at it. I understood this stuff before I heard this but it changed the way I saw species anyways.

I hope it wasn't boringly basic for many of you either :)

John Harshman · 22 January 2011

Jim Thomerson said: 20th century biologist me has been thinking about the Phylogenetic Species Concept. Is this a reasonable statement of the concept? A species originates at a speciation event and exists through time until it becomes extinct.
No, that's not the phylogenetic species concept. I'm not sure it has a name. Perhaps the lineage species concept? It resembles what Willi Hennig used, though I believe he also thought it was an artificial convention. Now, phylogenetic species can have varying definitions, but one definition is as the minimum diagnosable unit. If a population has at least one fixed difference from another population, we call it a species. In practice, this means that detectably different allopatric populations are called species. And that's one way to think about it.

Robert Byers · 22 January 2011

What's the problem?
AIG is right on.
They are simply from migration from Babel.
I would add that this all once again demands presumptions about genetics.
The Asian type would be a adaptation that happened after leaving the rest of mankind. so it follows the earliest remains would show likeness to other peoples that remained rather pure upon settlement. I mean New Guinea folk have been there from the beginning, and without much later people joining them, and so their genetics would be least changed from the original. The Asian in between simply from adaptation needs or interbreeding changed more in genetics.
These remains are simply early Asians before genetic variety. They are not directly related to New Guineas but the same original people expanding everywhere.
Creationism can always give a better answer because its based on better foundations.

mrg · 22 January 2011

Robert Byers said:[SNIP]
It's Saturday morning. The cartoons are starting early.

DS · 22 January 2011

Henry J said: Identifying the exact point at which a species branches off from another is a bit like identifying the precise boundary between a tree branch and the tree trunk (or its parent branch). Henry
Exactly. The distinction will always be a somewhat arbitrary point along a continuum. But it's even worse than that, because sometimes parts can swap after the branch point, maybe even long after the branch point. Still, genetic divergence seems to be the best way to decide these issues. It allows for the determination of reproductive isolation, time of divergence and even introgressive hybridization. It might still be somewhat arbitrary, but at least it is a real measure of something important. Creationists, on the other hand, still seem to be stuck back in the time of Linnaeus. They still seem to be trying to decide how many distinct, separate and unchanging species there are, while reality moves on without them.

Jim Foley · 22 January 2011

Robert Byers said: What's the problem? AIG is right on. They are simply from migration from Babel. I would add that this all once again demands presumptions about genetics. The Asian type would be a adaptation that happened after leaving the rest of mankind. so it follows the earliest remains would show likeness to other peoples that remained rather pure upon settlement. [etc., etc., etc.]
Dunning and Kruger strike again!

John Harshman · 22 January 2011

Henry J said: Still, genetic divergence seems to be the best way to decide these issues. It allows for the determination of reproductive isolation, time of divergence and even introgressive hybridization. It might still be somewhat arbitrary, but at least it is a real measure of something important.
Dang it, no. Genetic divergence doesn't allow for the determination of reproductive isolation. It allows for the determination of lack of gene flow. But lack of gene flow can have at least two causes: speciation or geographic isolation. In most species concepts, the latter doesn't count as speciation. Yes, it's a real measure of something important, but it isn't a measure of speciation. (Genetic divergence in sympatry is quite good evidence of separate species status. But divergence in allopatry isn't, and you can't infer the reverse, i.e. that lack of divergence means they're one species.)

stevaroni · 22 January 2011

DS said: Exactly. The distinction will always be a somewhat arbitrary point along a continuum. But it's even worse than that, because sometimes parts can swap after the branch point, maybe even long after the branch point.
Branches can also be forced to grow back together by the application of outside forces

harold · 22 January 2011

The concept of species is useful but also problematic, not least of all because a clear cut definition is elusive http://en.wikipedia.org/wiki/Species_problem.

I'll put aside the difficulty of applying the concept to prokaryotes (which are a massive part of the biosphere). Even in diploid/polyploid eukaryotes capable of meiosis, the concept is tricky.

At the current time, it is the human practice to subdivide the biosphere very finely into species.

This is tricky for a number of reasons.

First of all, many species have remarkably different free-living morphology at various times during development. Biologists aren't likely to be fooled by that any more but it was a problem historically.

Second of all, we define species very finely, and on the basis of behavior and morphology. We then run into the problem that some species, as we define them, interbreed with other species.

The biasing factor is that the implied definition of species is "any biological population that is sufficiently unique in some aspect of interest to humans to give it a separate name".

Also - as I noted back in the Neanderthal threads, any interbreeding between humans and other "hominid species" has strong implications. A low percentage of "Neanderthal genes" could mean limited interbreeding between populations of equal size - or it could mean complete assimilation of a much smaller Neanderthal population into a much larger H. sapiens population. Or it could mean something in between.

Jim Thomerson · 22 January 2011

I know the idea that a population with a single unique characteristic is seen as a species as the "Autapomorphic Species Concept". I don't think it is the same as the "Phylogenetic Species Concept", but maybe it is.

When I first studied what is now known as Gnatolebias zonatus and G. hoignei, the question was whether they were ecophenotypes of a variable species; the former in savanna pools and the latter in forest pools. At that time they had not been taken together. I found a consistent difference of in one of the fin ray counts, which made me think they were two species (Autapomorphy, I suppose). I did hybridization experiments and they wouldn't even try under conditions where both would spawn on a daily basis. So I described G. hoignei as new. Later we found their chromosomes were so different that production a a viable hybrid was unlikely, and their breeding behavior in nature was quite different. We also found them coexisting in an area cleared for agriculture, for at least several years, with no sign of interbreeding. DNA analysis has them as sister species.

DS · 22 January 2011

stevaroni said:
DS said: Exactly. The distinction will always be a somewhat arbitrary point along a continuum. But it's even worse than that, because sometimes parts can swap after the branch point, maybe even long after the branch point.
Branches can also be forced to grow back together by the application of outside forces
Right, but that doesn't mean they never existed. They can also grow around fences and other stuff as well. Living things tend to do messy stuff like that, given enough time.

RBH · 22 January 2011

Leszek said: Lets imagine we are standing face to face with a modern chimpanzee. We hold that hand of our mother and her other hand is holding the hand of her mother. So on it goes down the generations. Opposite us the chimp is doing the same. With an average separation of 1 meter (just over a yard for you non metric types) a line would stretch from Windsor (Canadian side of the boarder from Detroit) down the line until at around Toronto something interesting would happen. The Chimp line and your line would be holding the opposite hands of the same mother. If you stretch these two lines out to one continuous line, there is no point at which you can say everyone on this side is Human and everyone on that side is Chimp without breaking up a mother and child. It is only easy to identify one or the other because all the ancestors are already dead. The concept of species works really well in those situations.
A similar way to think of it is this: By any definition of "species" that I know of, every generation of a lineage is of the same species as its immediate parent generation, yet at the end of N thousands of generations one can have a population that is of a different species from the distant ancestral generation of N thousands of generations ago. Or as the man said, I can't say exactly when day ends and night begins, but I sure as hell can tell the difference between them. :)

RBH · 22 January 2011

By the way, John Wilkins (of Talkorigins and Evolving Thoughts fame) has the most current analysis of species concepts in Species: A history of the idea, which is coming out soon in paperback. Here is an overview post he wrote not long ago. From it:
It’s an old question in biology: what is a species? Many answers have been given over the years – I counted 26 in play, and recently a new one, the “polyphasic” concept (basically a consilience of many lines of evidence) has been introduced in bacterial and other microbial contexts, and which may apply to macrobial species too.

DS · 22 January 2011

RBH said: Or as the man said, I can't say exactly when day ends and night begins, but I sure as hell can tell the difference between them. :)
Thanks RBH. That's a good one. I'm going to use that.

Epimetheus · 22 January 2011

"Though no man can draw a stroke between the confines of night and day, yet light and darkness are upon the whole tolerably distinguishable." -- Edmund Burke

Jim Thomerson · 22 January 2011

Here is a scenario for you. A population of the species which is the last common ancestor of humans and chimps undergoes some sort of event which separates it into two allopatic populations. After a while, due to genetic drift, differential selection pressures, whatever, a speciation event has occurred. Now there are two species, one the first common ancestor of humans, and the other, the first common ancestor of chimps. The last common ancestor of humans and chimps is now extinct. That's my story and I'm stickin' to it.

veritas36 · 22 January 2011

A snippet of conversation

Guy who is annoyed by creationism, "How did all the animals fit on the ark?"

Student at Liberty Baptist, seriously: They didn't have to take each variety, not a pair of polar bears, brown bears, grizzly pairs, black bears. Just one pair of bears was enough."

I regret to say I don't know what the explanation for how all the different kinds of bears reappeared.

veritas36 · 22 January 2011

Neanderthal & Denisovan contributions to some humans' genomes are ~5% despite a long breeding interval after they became extinct. Assuming the interbreeding occurred in stable mating pairs, this suggests that Neanderthals & Denisovans could talk. Or that modern humans could not, at the time of interbreeding.

I suggest that human societies would disfavor matings with beings that can't talk. Tribal support is evolutionary significant, and talking is distinctly useful as well.

mrg · 22 January 2011

veritas36 said: I regret to say I don't know what the explanation for how all the different kinds of bears reappeared.
That's where "created kinds" comes into play. All the Ark had to do was carry one pair of each "kind". On recovery, they went through a phase of hyperevolution (yes, creobots will embrace evolution, if only for as long as it takes them to get out of a tight spot). I regret to say that I DO know the explanation. Fortunately, I can generally keep suchlike segregated on the exterior side of my baloney filter.

Mike Elzinga · 22 January 2011

Are the iron and oxygen in FeO, Fe3O4, and Fe2O3 “different species” of iron and oxygen?

The crystalline structures of bulk quantities of these compounds are different. Are these different species?

Does that mean that the crystals have different gene pools?

What happens when one form evolves into another given the appropriate environment? Is that natural selection producing different species?

How about the many forms of even a single atom carbon? There is graphite, diamond, fullerenes, graphene, etc.. Between the carbon and its many collective forms, which are the “genes” and which are the “species?”

What about the icicles hanging from eaves? As the days go by, they come and go and branch. Where is the common ancestor of two different icicles? Where in a bifurcation do two icicles become different icicles?

Start with any huge collections of atoms and molecules all doing their thing in whatever environment is current. If some of these collections start going off in a different direction than others, how is that any different from the issues involved in speciation?

The only thing we can say they are different when there are enough accumulated differences to allow us to distinguish within a given resolution.

Tiny differences in patterns can be perceived by rapidly switching back and forth between them; otherwise they are indistinguishable.

This entire discussion over speciation is only a problem because of the historical tradition of classifying and cataloguing things that were clearly different (at least in appearance). As technology gets better, so does the resolution; but below that resolution, we can’t distinguish.

david winter · 22 January 2011

John Harshman said:
Jim Thomerson said: 20th century biologist me has been thinking about the Phylogenetic Species Concept.
No, that's not the phylogenetic species concept. I'm not sure it has a name. Perhaps the lineage species concept? way to think about it.
Either the "general lineage species concept" or the "evolutionary species concept". Part of the problem of dealing with these terms,as you pointed out, is that we want 'species' to mean the same thing when we look at modern species as we look at fossil species. It's possible that "lineage species" could go through multiple distinct morphlogical forms over time and get given different names by paleontologists over the course of its evolution

Mike Elzinga · 22 January 2011

mrg said:
veritas36 said: I regret to say I don't know what the explanation for how all the different kinds of bears reappeared.
That's where "created kinds" comes into play. All the Ark had to do was carry one pair of each "kind". On recovery, they went through a phase of hyperevolution (yes, creobots will embrace evolution, if only for as long as it takes them to get out of a tight spot). I regret to say that I DO know the explanation. Fortunately, I can generally keep suchlike segregated on the exterior side of my baloney filter.
I wonder how many different species of religion they would say there were. Given their penchant for microscopic hermeneutics, exegesis, etymology, and general word gaming, their ability to distinguish between “Christians” and “non-Christians” should be sharply honed.

mrg · 22 January 2011

Mike Elzinga said: I wonder how many different species of religion they would say there were.
There are clear distinctions. "What's the difference between Catholics and Mormons?" "Mormons greet each other at the liquor store." I've worked with a number of Mormons and they usually like that joke.

Paul Burnett · 22 January 2011

veritas36 quoted: Student at Liberty Baptist, seriously: "They didn't have to take each variety (on Noah's Ark), not a pair of polar bears, brown bears, grizzly pairs, black bears. Just one pair of bears was enough."
I've had creationists patiently explain to me that there was one "kind" of beetle and one kind of "germ" and so on. When I explain that that means the subsequent hyperevolution meant some creatures would have to change species several times in their lifetimes, they would get all upset.

John Harshman · 22 January 2011

Jim Thomerson said: Here is a scenario for you. A population of the species which is the last common ancestor of humans and chimps undergoes some sort of event which separates it into two allopatic populations. After a while, due to genetic drift, differential selection pressures, whatever, a speciation event has occurred. Now there are two species, one the first common ancestor of humans, and the other, the first common ancestor of chimps. The last common ancestor of humans and chimps is now extinct. That's my story and I'm stickin' to it.
That was Willi Hennig's solution. Like I said, I believe he realized it was a convention, nothing more. It makes a nice dividing line on a phylogenetic tree, but of course we know that in reality there was no such line. It's a form of the chronospecies concept, which is another of the several bad ways to extend species concepts into evolutionary time.

John Harshman · 22 January 2011

Mike Elzinga said: Are the iron and oxygen in FeO, Fe3O4, and Fe2O3 “different species” of iron and oxygen?
Well, the irons are. That is in fact what chemists call them. But it may not surprise you to learn that the same word can have different meanings in different contexts. I'm not sure of the relevance of bringing non-organisms into it. But "species" does have a real meaning in biology. I don't see a reason to dispense with it

Les Lane · 22 January 2011

But why do you want to introduce a term for a quality that you cannot define?
The term "species" is an historical relic of the time when species were defined by morphology and believed to be separate and distinct.

DS · 22 January 2011

Les Lane said:
But why do you want to introduce a term for a quality that you cannot define?
The term "species" is an historical relic of the time when species were defined by morphology and believed to be separate and distinct.
Exactly. If you can't define species, that's a problem for creationism, not evolution.

Mike Elzinga · 22 January 2011

John Harshman said:
Mike Elzinga said: Are the iron and oxygen in FeO, Fe3O4, and Fe2O3 “different species” of iron and oxygen?
Well, the irons are. That is in fact what chemists call them. But it may not surprise you to learn that the same word can have different meanings in different contexts. I'm not sure of the relevance of bringing non-organisms into it. But "species" does have a real meaning in biology. I don't see a reason to dispense with it
I was merely pointing out that condensed matter systems, no matter what their level of complexity, can and do branch. These may be something as distinct as different compounds – which can change into other forms under suitable environmental conditions – or they can be extremely complex organic systems involving molecules with different chirality, or replicating systems producing approximate surrogates that survive and reproduce differentially. Any of these systems can have bifurcation ranges below the technological limits of resolution. The word species may have a certain historical context in biology where it has been given distinct meanings depending on the level of complexity of the organisms involved. But the issue of making distinctions among species and where they branch to become “separate” species remains the same at some fundamental level whether or not these systems reproduce surrogates that branch or whether it is a single system that continues on and branches to adjust to its environment. In casting objects from molten metals or some other substance that solidifies in a mold, we don’t say that the cast metal has evolved. But with successive castings of the same metal, we might see design modifications in the casting, or we may see completely different objects. The terms that describe those differences apply to those higher levels provided that these higher levels are conceptually distinguishable from the constituents in its substrate. But when zooming in at the molecular level, how is one to discuss the differences between successive designs of a casting? There is a crossover point below which it is not possible to learn from the molecular level what will appear at the macroscopic level. This is not mere semantics; it is one of the apparent paradoxes of condensed matter systems. It does not appear to be possible to predict the characteristics of a macroscopic system from detailed knowledge of its atomic and molecular constituents. The reason is because collective emergent properties come so quickly and in such quantity that one simply doesn’t have enough knowledge at any given level to predict where things will go at levels above. One has to have some knowledge of those emergent properties in order to make better use of the detailed knowledge of the constituents in predicting how a system will evolve. The point of all this – and this includes biology – is that there will very likely always be a fundamental lower limit below which it will not be possible to distinguish among species; but as RBH has already pointed out, at some level we will know it when we see it. That should not cause anyone any concern in any rational world. But in a world containing the political shenanigans of creationists, we see this mud wrestling over speciation far more than we need to.

Jim Thomerson · 22 January 2011

Most species known today are defined on morphology and are thought to be separate and distinct. It's a grand tradition.

Mike Elzinga · 22 January 2011

Jim Thomerson said: Most species known today are defined on morphology and are thought to be separate and distinct. It's a grand tradition.
It’s a fine tradition. And it can be adjusted to whatever level everyone agrees they see significant differences. As to ability to interbreed in a shared gene pool, somewhere down at the molecular level might be adequate; provided that enough knowledge of the levels above that insure the difference really does take place down there. I would suspect that the general rule should be that a species difference can be defined at any level that has been established as the one that makes the difference. That probably is most likely established experimentally with some input from theoretical calculations.

Henry J · 23 January 2011

Exactly. If you can’t define species, that’s a problem for creationism, not evolution.

It's also a problem for anybody who wants a count of how many species are present in a region, an ecosystem, or the whole planet.

Hygaboo Andersen · 23 January 2011

Ron Bear said: Could someone please explain to me what "the Denisovans seem to have made about a 5% contribution to the genome of living Melanesians" means. I know I am doing something stupid, but I don't know what. We're 98 percent the same genome as Chimps, but Melanesians got 5 percent from Denisovans. Is that 5 percent of the 2 percent that makes us different from Chimps? It is obvious to me that the 98 percent and the 5 percent can't be a percentage of the same thing, but I'd like to understand what I am missing here. Thanks in advance. Ron
What I think the evolutionists mean by this is that modern Melanesians are descendant from a combination of Denisovans and monkey while other people are descendant from only monkeys.

Hygaboo Andersen · 23 January 2011

Jeremiah Tattersall said: I've talked with my local open air creationist preacher about the new finding about Denisovan (we have weekly coffee then screaming matches). After we got through all the BS about dating methods and sequencing his argument position was that of Magic Man done it. To quote: "If the fossils seem old it's because god made them seem that way. If the DNA seems like it was sequenced it's because god made it seem that way, not because it actually is that way". That's right. God, the great deceiver.
There are other possibilities too. They could be chickens with arthritis. They could be ancient statues or parts of them. Remember Nebraska Man? The evolutionists simply read what their religion tells them in every fossil--real or imagined. And they think Christians who see the Virgin Mary or Jesus in a tortilla is stupid. They are just as bad!

Joe Felsenstein · 23 January 2011

Hygaboo Andersen said:
Ron Bear said: Could someone please explain to me what "the Denisovans seem to have made about a 5% contribution to the genome of living Melanesians" means. I know I am doing something stupid, but I don't know what. We're 98 percent the same genome as Chimps, but Melanesians got 5 percent from Denisovans. ... [and so on, adequately answered by me earlier]
What I think the evolutionists mean by this is that modern Melanesians are descendant from a combination of Denisovans and monkey while other people are descendant from only monkeys.
A careful technical comment on this is: you haven't a clue. You need to learn a little paleoanthropology. Just a little would make a big difference. You are so far off that all I can do is to "lose it" and say that monkeys are descendants of Denisovans and creationists, crossed with trolls. (I will not further respond to Hygaboo because Hygaboo is just trolling and there is no sign that Hygaboo is even a serious creationist).

John Kwok · 23 January 2011

Robert Byers said: What's the problem? AIG is right on. They are simply from migration from Babel. I would add that this all once again demands presumptions about genetics. The Asian type would be a adaptation that happened after leaving the rest of mankind. so it follows the earliest remains would show likeness to other peoples that remained rather pure upon settlement. I mean New Guinea folk have been there from the beginning, and without much later people joining them, and so their genetics would be least changed from the original. The Asian in between simply from adaptation needs or interbreeding changed more in genetics. These remains are simply early Asians before genetic variety. They are not directly related to New Guineas but the same original people expanding everywhere. Creationism can always give a better answer because its based on better foundations.
Absolutely priceless, Booby. 8.5 for content, 9.2 for hilarity. Having dealt with an online stalker over on Facebook who has been obsessed with me and two well known writers - one a friend, the other an acquaintance of mine - over the past few days, I always know I can depend on you for some laughs!!!!

John Kwok · 23 January 2011

Les Lane said:
But why do you want to introduce a term for a quality that you cannot define?
The term "species" is an historical relic of the time when species were defined by morphology and believed to be separate and distinct.
No, as John Harshman has pointed out, this is still a valid definition of species, relied on by past, current and future generations (of which I have no doubt) of systematists.

Frank J · 23 January 2011

Exactly. If you can’t define species, that’s a problem for creationism, not evolution.

— DS
Thank you! Now if we can only get the public to understand that irony. Though "hard-to-define" is not the same as "not useful." The most obvious analogy is "branch." At the tips, it's easy to differentiate branches, but not so near the base. If anything it's more than an analogy, because life is arranged in a "tree." Yet that concept has not sunk in with most nonscientists. Creationism - when it's not actually conceding a ~4 billion year old "tree" as in the case of Behe and his "kind" - claims a "lawn" instead. While the "tree" may require changes that some people find "impossible," the "lawn" requires even far more radical changes, namely the independent origin of many lineages from nonliving matter, not just "major genome changes." Invoking "common design," whether or not warranted, does not in any way help the "lawn" hypothesis, especially the absurdly recent one that many, but not all, creationists prefer. But I suspect that most "lawn" peddlers know that, or else they'd devote more effort to get Behe and his "don't ask, don't tell" buddies on board.

DS · 23 January 2011

Hygaboo Andersen said: What I think the evolutionists mean by this is that modern Melanesians are descendant from a combination of Denisovans and monkey while other people are descendant from only monkeys.
DS said: Apparently, there was some interbreeding in the past of some Denisovans and some modern humans. Apparently some of these genes, which can be distinguished from the lineage they introgressed into due to the degree of genetic divergence, can still be found in some modern populations. That is where the 5% figure comes from. It doesn't have anything to do with the divergence of humans from chimps, since they last shared a common ancestor before the split of modern humans and Denisovans.
Asked and answered peekaboo. Move along.

John B. · 23 January 2011

In a nutshell, "they're all just humans".

Jim Thomerson · 23 January 2011

Looking at COPEIA, 2010 #4, I find descriptions of one new frog, and 12 new fish species. All are based on morphological differences from known species, and two also include comparison of allozyme patterns. There is an article on use of gill pore papillae morphology for identifying lamprey species, and a report of the first known natural hybridization event between largemouth and smallmouth black basses. All very interesting!

Ron Okimoto · 23 January 2011

It isn't just the 8 people on the ark that you have to worry about. We will eventually have whole genome sequences (from multiple individuals) of the Denisovans and Neandertals. They will have to try to reconcile how three such distinct populations could arise from one man and one woman made from the rib of the man. There isn't much genetic diversity among these two individuals especially since they were considered to be pinnacle of creation.

There are pretty much only two options. Either a huge amount of genetic variation was packed into these two individuals and segregated out, or an equally huge amount of genetic variation is due to mutation after the fall. There could be a combination of the two, but even the amount of each that had to occur in the combination is mind boggling if you only have a couple thousand years to generate it.

The first one doesn't stand up to scrutiny because we will have whole genome sequences and there will be very little chance that enough recombination events occurred within the possible four haplotypes that existed in the first two individuals to generate the distinct lineages.

The second fails because there is too little time for the mutations to accumulate. It isn't just absurdly high mutation rates that are required, but it is likely that the Denisovan and Neandertal genomes will show evidence of as high amounts of purifying selection as is found in modern human genomes. For a coding sequence, by chance, the number of replacement substitutions (mutations that change the amino acid sequence of the encoded protein)should outnumber silent substitutions by around 2:1, but we find that the number of silent mutations outnumber the replacement substitutions by 2:1 or more. This means that a whole lot of replacement substitutions have been removed by selection. These replacements had a large enough negative effect on the population that they were selected out of the gene pool. As certain creationist have pointed out there is a cost to selection. Real science has several orders of magnitude more time for this selection to have occurred than just a couple thousand years.

There is one possible out. The Nephlim of the Bible were supposed to be the sons of the gods that had bred with humans. Since we have no god DNA to compare we can't say how different the various gods were in the DNA that they transmitted to the human stock. 5 backcrosses to any single god lineage would produce genomes over 95% of that god lineage. So you could create distinct populations in this way. We would then have to figure out if the 5% of our DNA (for those or us that are of relatively recent non African origin) came from hanky panky with a god like Baal or his Neandertal offspring before we migrated out of the middle east, and took over the rest of the world after the flood. Apollo or Thor could have left their mark on the New Guineans during a tropical vacation.

Maybe the Neandertals or the Denisovans were the true purely human descendants of Adam and Eve. It should be a hoot to watch the YECers try to use the sequence data to try and sort that one out.

Scott F. · 23 January 2011

Mike Elzinga said: I was merely pointing out that condensed matter systems, no matter what their level of complexity, can and do branch.
Perhaps a concrete example of what you are describing might help. For example, looking just at a single carbon atom and its bonds, you can't tell what assemblage it belongs to. Step out one level of complexity, and you see it is part of a hexagonal ring of carbon atoms. But you still can't tell what assemblage it belongs to. Step out another level, and you see that this carbon ring is part of a sheet of interconnected carbon rings. Only at the next level out can you tell if this pattern is part of graphite or of a nano-tube.

Paul Burnett · 23 January 2011

Ron Okimoto said: It should be a hoot to watch the YECers try to use the sequence data to try and sort that one out.
As if there are any YECs who could "try to use the sequence data" - 99.99+ percent of YECs don't have a clue what the term means: Genome sequencing isn't mentioned in Genesis, so it can't exist in their universe.

Mike Elzinga · 23 January 2011

Scott F. said:
Mike Elzinga said: I was merely pointing out that condensed matter systems, no matter what their level of complexity, can and do branch.
Perhaps a concrete example of what you are describing might help. For example, looking just at a single carbon atom and its bonds, you can't tell what assemblage it belongs to. Step out one level of complexity, and you see it is part of a hexagonal ring of carbon atoms. But you still can't tell what assemblage it belongs to. Step out another level, and you see that this carbon ring is part of a sheet of interconnected carbon rings. Only at the next level out can you tell if this pattern is part of graphite or of a nano-tube.
Not only is this a nice example, it illustrates exactly the problem of predicting higher levels of complexity from lower level properties in any given system. Because, as it turns out, it has been possible with carbon to actually predict which kinds of structures are relatively stable with carbon atoms. Most of the forms of carbon were predicted before they were actually observed. But that doesn’t mean that any particular evolutionary process to a particular form can be predicted. To do this requires far more detailed knowledge of every contingency in the processes leading to the growth of a structure than could possibly be made available. And these contingencies depend on what has already been produced at any given stage in a process. The only thing that can be done in this case it to use information about the binding energies of these structures to estimate their stability, and from that to estimate the probability that a given structure would be produced in a given process of growing these structures. From this one can try to pick the process that will produce the highest percentages of the desired structure.

M.W. · 23 January 2011

If a person is brought up to hear and read the word of God then it is only natural that they would believe that the creation was by God. John ch 1 vs 1-3 In the beginning was the word, and the word was with God ……..then in vs 4 In him was life and the life was the light of man. It is the beginning of an explanation of the creation, Genesis ch 1 vs 1 In the beginning God created the heavens and the Earth. Genesis ch 2 vs 1-3 ……God rested from all his work which he had done in creation, then in vs 4 These are the generations of the heavens and the Earth. It doesn't go into the chemistry that is there for adventure to analyze. Jesus couldn't comment on evolution as Darwin wasn't around at that time. Jesus never implied that he was anything but the son of man through out the history of the Earth, Philippians ch 2 v 6-7….being born in the likeness of men. In Genesis ch 1 vs 26-27 Then God said let us make man in our image in our likeness.... Then Genesis ch 3 vs 22 ......the man has become like one of us.
Jesus left us with a new covenant John ch 13 vs 34 and that was to love one another, that is one of the things that not only humans can do. What direction does evolution offer or Darwin, only that the animal kingdom naturally selects, but to what end, I feel there is more to life than that. John ch 15 vs 5. There is a purpose for the branch and that is that it goes in the right direction.

Mike Elzinga · 23 January 2011

M.W. said: What direction does evolution offer or Darwin, only that the animal kingdom naturally selects, but to what end, I feel there is more to life than that. John ch 15 vs 5. There is a purpose for the branch and that is that it goes in the right direction.
But the “right direction” suggests, after the fact, that humans were the goal of evolution. This is a well-understood fallacy called the Lottery Winner’s Fallacy. It conflates the probability that a particular winner has won with the probability that someone has won.

Flint · 23 January 2011

It conflates the probability that a particular winner has won with the probability that someone has won.

And is even more suspect because the "winners" of this particular lottery got to postdict the rules that made them the winners. Of course, what we should recognize is that some games of chance ARE fixed, and cheating is not unknown. So if we wish to believe that some magical entity rigged the game to engineer a predetermined outcome, why not?

Wheels · 23 January 2011

I'd just like to go OT for a moment and thank the commenters for always giving me interesting diversions. I never thought that a post about the genetic differences between different species of humans would have me reading about ducks on Wikipedia for an hour.

Frank J · 23 January 2011

Paul Burnett said:
Ron Okimoto said: It should be a hoot to watch the YECers try to use the sequence data to try and sort that one out.
As if there are any YECs who could "try to use the sequence data" - 99.99+ percent of YECs don't have a clue what the term means: Genome sequencing isn't mentioned in Genesis, so it can't exist in their universe.
Certainly 99.99+% of creationists-on-the-street (YEC or otherwise) don't. But YEC, and especially ID, activists know just enough to fool millions of non-creationists-on-the-street. If only for the simple reason that refutations of those misleading sound bites are less likely to be heard, let alone understood and remembered.

Mike Elzinga · 23 January 2011

Flint said: And is even more suspect because the "winners" of this particular lottery got to postdict the rules that made them the winners.
Yeah; that’s the game, isn’t it? And you make it seem impossible by stringing those detailed rules by the millions in succession and then multiplying all the probabilities of each step together. Ergo; deity done it!

Scott F · 23 January 2011

M.W. said: It [the Bible] doesn't go into the chemistry that is there for adventure to analyze. Jesus couldn't comment on evolution as Darwin wasn't around at that time. Jesus never implied that he was anything but the son of man through out the history of the Earth,...
Yes, but the chemistry was still there, as was evolution. Chemistry didn't magically start with Tapputi, and evolution didn't magically start with Darwin. When God created the world, he created both chemistry and evolution at the same time. If Jesus had any divine omniscience, he would have known about both.

fnxtr · 23 January 2011

M.W. said: What direction does evolution offer or Darwin, only that the animal kingdom naturally selects, but to what end, I feel there is more to life than that.
Probably true, M.W., but also not really the point. If by "life" you mean some existential "meaning" to human history, then you are of course correct, neither Darwin, nor the fact, nor theory that explains the fact of evolution, ever intended to address that issue. If by "direction" you mean "pre-ordained goal/target", well, evolution seems not to have one. As has been mentioned before, it's not 'survival of the fittest', it's 'reproduction of the fit enough', also coarsely described as 'live long enough to get laid'. But these are just the bare facts. Beyond that, life is what we make it. That's what the gift of free will is all about, isn't it? What's "more to life" is ours to decide.

SAWells · 24 January 2011

Hmm. Species form continua if you're looking back in time but are usually discrete when taking a slice at any moment in time, just as tree branches all connect if you trace them back to the trunk but are discrete if you apply a hedge trimmer. Sometimes we see a case where branching is ongoing.

And our usual quote from Burke: "Though no man can draw a stroke between the confines of day and night, yet light and darkness are upon the whole tolerably distinguishable."

Maybe "Though no man can draw a stroke between the confines of dog and wolf, yet sharks and labradors are upon the whole tolerably distinguishable."

John Kwok · 24 January 2011

david winter said:
John Harshman said:
Jim Thomerson said: 20th century biologist me has been thinking about the Phylogenetic Species Concept.
No, that's not the phylogenetic species concept. I'm not sure it has a name. Perhaps the lineage species concept? way to think about it.
Either the "general lineage species concept" or the "evolutionary species concept". Part of the problem of dealing with these terms,as you pointed out, is that we want 'species' to mean the same thing when we look at modern species as we look at fossil species. It's possible that "lineage species" could go through multiple distinct morphlogical forms over time and get given different names by paleontologists over the course of its evolution
Actually the correct term would be the "chronospecies concept" as noted here: http://evolution.berkeley.edu/evosite/evo101/VA1BioSpeciesConcept.shtml I would concur with your observation david if most lineages did indeed show "phyletic gradualism", but, on the contrary, there seems to be widespread data in support of rapidly brief divergence at the onset and long-term morphological stasis, which is what one would expect from "punctuated equilibrium".

eric · 24 January 2011

veritas36 said: I suggest that human societies would disfavor matings with beings that can't talk. Tribal support is evolutionary significant, and talking is distinctly useful as well.
You've never traveled to a foreign country and met a hot woman that didn't speak your language? Compared to matings with people you can speak to, its probably relatively rare. But given human proclivities it would be crazy to think it didn't happen at all. It happens now, after all.
M.W. said: What direction does evolution offer or Darwin, only that the animal kingdom naturally selects, but to what end, I feel there is more to life than that.
And I felt that life should be fair, and was deeply troubled when I found out it isn't - back when I was about eight. As an adult, I got over it and learned to accept the fact that life does not run according to the rules I feel it ought to run by.

fusilier · 24 January 2011

M.W. said: If a person is brought up to hear and read the word of God then it is only natural that they would believe that the creation was by God. John ch 1 vs 1-3 In the beginning was the word, and the word was with God ……..then in vs 4 In him was life and the life was the light of man. It is the beginning of an explanation of the creation, Genesis ch 1 vs 1 In the beginning God created the heavens and the Earth. Genesis ch 2 vs 1-3 ……God rested from all his work which he had done in creation, then in vs 4 These are the generations of the heavens and the Earth. It doesn't go into the chemistry that is there for adventure to analyze. Jesus couldn't comment on evolution as Darwin wasn't around at that time. Jesus never implied that he was anything but the son of man through out the history of the Earth, Philippians ch 2 v 6-7….being born in the likeness of men. In Genesis ch 1 vs 26-27 Then God said let us make man in our image in our likeness.... Then Genesis ch 3 vs 22 ......the man has become like one of us. Jesus left us with a new covenant John ch 13 vs 34 and that was to love one another, that is one of the things that not only humans can do. What direction does evolution offer or Darwin, only that the animal kingdom naturally selects, but to what end, I feel there is more to life than that. John ch 15 vs 5. There is a purpose for the branch and that is that it goes in the right direction.
MW, since you like the Gospel according to John, how about John 6:46-56? Do you take that literally? The Pope does, and Benedict XVI, John Paul II, and Pius XII have all said there are no problems with the biological theory of evolution, specifically with regard to the evolution of the human body. fusilier James 2:24

DS · 24 January 2011

Scott F said:
M.W. said: It [the Bible] doesn't go into the chemistry that is there for adventure to analyze. Jesus couldn't comment on evolution as Darwin wasn't around at that time. Jesus never implied that he was anything but the son of man through out the history of the Earth,...
Yes, but the chemistry was still there, as was evolution. Chemistry didn't magically start with Tapputi, and evolution didn't magically start with Darwin. When God created the world, he created both chemistry and evolution at the same time. If Jesus had any divine omniscience, he would have known about both.
Indeed. If Jesus would have just explained how evolution worked, it would be a lot easier to accept his claim to being god. After all, presumably the whole thing was his idea in the first place. But then again, he didn't bother to explain germ theory either now did he?

Kim · 24 January 2011

(Indeed, given the original definition of species as referring to organisms that could interbreed successfully, treating them as separate species doesn’t make sense. However, that definition is no longer observed.)
The interbreeding criteria is used for extant species, not for the continuous line of gradually changing species. That is why the biological species concept is only one of many species concept and using a phylogenetic species concept would make more sense here.

John Harshman · 24 January 2011

John Kwok said: Actually the correct term would be the "chronospecies concept"
No, a chronospecies is an arbitrary division of a lineage. The concept being discussed was an entire branch of the tree, from speciation to extinction (and counting new speciation as extinction of the ancestor). That's quite different.
I would concur with your observation david if most lineages did indeed show "phyletic gradualism", but, on the contrary, there seems to be widespread data in support of rapidly brief divergence at the onset and long-term morphological stasis, which is what one would expect from "punctuated equilibrium".
The problem with PE is that it assumes biological species and morphospecies are the same, for which there is no evidence. All the paleontological data can show, at most, is that morphological evolution is characterized by episodes of rapid and slow change. But it can't show that the rapid episodes are coincident with speciation in the BSC sense, i.e. branching events. Or, to put it in Gouldian terms, we can't distinguish between punctuated phylogenesis and punctuated anagenesis.

Jim Thomerson · 24 January 2011

I should mention that my BS and MS were in geology and I did a study of Cretaceous marine micro fossils for my MS thesis.

I did a little reading up on the Phylogenetic Species Concept. It is seen in different lights by different folks (Isn't that amazing?) But the theme of monophyly and uniqueness (autapomprphy) seem widespread. I don't have any problem with that.

My favorite species concept is the "Sophisticated Biological Species Concept" which is biological species as I understand it applied a particular case in the real world. However, the most widely used species concept is the morphospecies concept. The large majority of modern day animals were described as morphospecies. I described Rivulus corpulentis based on a jar of preserved fish on the shelf at the California Academy of Sciences. Clearly a morphospecies, but I am confident that should it be examined in nature, it will be found to be a biological species.

So I regard morphospecies as a stand in for biological species. And, generally, when we are able to study the situation further, do hybrid experiments, look at syntopic relatives, etc. the morphospecies turns into the biological species it represents.

I don't see a difference in kind between modern and fossil morphospecies, just a difference in how easy it is to decide if they are actually biological species.

Michael Roberts · 24 January 2011

Scott F said:
M.W. said: It [the Bible] doesn't go into the chemistry that is there for adventure to analyze. Jesus couldn't comment on evolution as Darwin wasn't around at that time. Jesus never implied that he was anything but the son of man through out the history of the Earth,...
Yes, but the chemistry was still there, as was evolution. Chemistry didn't magically start with Tapputi, and evolution didn't magically start with Darwin. When God created the world, he created both chemistry and evolution at the same time. If Jesus had any divine omniscience, he would have known about both.
Why should Jesus have any divine omniscience. after all he was born a little baby. Further Philippians 2 makes it clear that any view of the Incarnation makes it clear that Jesus cast aside his god-attributes during his earthly life. Hence he could have been a flat-earther and that does not affect my faith

John Kwok · 24 January 2011

John Harshman said:
John Kwok said: Actually the correct term would be the "chronospecies concept"
No, a chronospecies is an arbitrary division of a lineage. The concept being discussed was an entire branch of the tree, from speciation to extinction (and counting new speciation as extinction of the ancestor). That's quite different.
I would concur with your observation david if most lineages did indeed show "phyletic gradualism", but, on the contrary, there seems to be widespread data in support of rapidly brief divergence at the onset and long-term morphological stasis, which is what one would expect from "punctuated equilibrium".
The problem with PE is that it assumes biological species and morphospecies are the same, for which there is no evidence. All the paleontological data can show, at most, is that morphological evolution is characterized by episodes of rapid and slow change. But it can't show that the rapid episodes are coincident with speciation in the BSC sense, i.e. branching events. Or, to put it in Gouldian terms, we can't distinguish between punctuated phylogenesis and punctuated anagenesis.
Sorry John, as a former paleobiologist, I respectfully beg to differ. Starting with Niles Eldredge's work on phacopid trilobites, we do see instances of branching events in the metazoan fossil record that would substantiate "speciation" events yielding to long-term morphological stasis of a given species (which I will concede that if one were to operate strictly on a biological species concept, there is no way to distinguish into a morphospecies any instances of sympatric speciation). For "punctuated anagenesis" to occur, the best examples are within the protistan fossil record, especially amongst the radiolarians and foraminifera.

John Harshman · 24 January 2011

John Kwok said: Sorry John, as a former paleobiologist, I respectfully beg to differ. Starting with Niles Eldredge's work on phacopid trilobites, we do see instances of branching events in the metazoan fossil record that would substantiate "speciation" events yielding to long-term morphological stasis of a given species (which I will concede that if one were to operate strictly on a biological species concept, there is no way to distinguish into a morphospecies any instances of sympatric speciation). For "punctuated anagenesis" to occur, the best examples are within the protistan fossil record, especially amongst the radiolarians and foraminifera.
Sorry, John. As a current systematist, I also respectfully beg to differ. We have no way of knowing if those were branching events. If we find two disparate morhpologies in the same locality, we can often suppose that they are separate species (with caveats for sexual and other dimorphism, ecotypes, preservational artifacts), but the appearance of two such morphotypes can often be explained by migration. Or, if geographic sampling is good, as with the Eldredge Phacops instance, we can't tell if there were already two morphologically similar species before one of them experienced punctuated anagenesis. And cryptic/sibling species are very common, and would be even more common if we had only the limited information for extant species that can be gleaned from fossils. I'm certainly not the first to point this out.

harold · 24 January 2011

eric -
And I felt that life should be fair, and was deeply troubled when I found out it isn’t - back when I was about eight.
I was raised in a traditional religion that I had no problem with. I was aware that the world did not operate by the rules I wished it would, which actually overlapped a great deal with the rules of being a "Christian" at that time, at a fairly early age. (Times have changed massively. Back then doing things like gunning down doctors would have been seen as "un-Christian".) I was always very interested in animals, and that may have been part of what raised my awareness that things were "imperfect" from a human perspective. And "the fall" just isn't a very intellectually satisfying The theory of evolution made life make sense to me. I more or less got the idea before taking any formal treatment of evolution. I had a disrupted high school career, and when I actually managed to jam my way into university despite that, I changed majors from accounting to biology and ended up taking molecular biology, cell biology, and biochemistry before getting a formal evolution course. As I was noticing that life has common descent and evolves and adapts in a way that doesn't need to be explained by magic, I often had a "why didn't I think of this before?" feeling. I also notice that many of the problems that humans have with the theory of evolution, putting aside the pure right wing cult follower driven denial of contemporary America, come from projection. Most of the biosphere consists of plants and microbes, and there's not much reason to think that most animals are ridden with existential angst or ennui. We can philosophize, perhaps aided by mildly psychoactive substances, about the fact that evolution produced a brain/mind that now contemplates evolution, but acorns almost certainly don't give a damn whether or not they grow up to make other acorns. The fact that we can perceive that a process of selection is often part of the evolution of life on earth is not a reason to think that we, or any other living beings, should find life "pointless", meaningless", etc.

mrg · 24 January 2011

harold said: The fact that we can perceive that a process of selection is often part of the evolution of life on earth is not a reason to think that we, or any other living beings, should find life "pointless", meaningless", etc.
From Terry Pratchett's THE HOGFATHER, Death talking to his granddaughter Susan (don't ask for details on that relationship):
Death: Humans need fantasy to *be* human. To be the place where the falling angel meets the rising ape. Susan: With tooth fairies? Hogfathers? [The Diskworld analog of Santa Claus] Death: Yes. As practice, you have to start out learning to believe the little lies. Susan: So we can believe the big ones? Death: Yes. Justice, mercy, duty. That sort of thing. Susan: They're not the same at all. Death: You think so? Then take the universe and grind it down to the finest powder, and sieve it through the finest sieve, and then show me one atom of justice, one molecule of mercy. And yet, you try to act as if there is some ideal order in the world. As if there is some, some rightness in the universe, by which it may be judged. Susan: But people have got to believe that, or what's the point? Death: You need to believe in things that aren't true. How else can they become?

John Kwok · 24 January 2011

John Harshman said:
John Kwok said: Sorry John, as a former paleobiologist, I respectfully beg to differ. Starting with Niles Eldredge's work on phacopid trilobites, we do see instances of branching events in the metazoan fossil record that would substantiate "speciation" events yielding to long-term morphological stasis of a given species (which I will concede that if one were to operate strictly on a biological species concept, there is no way to distinguish into a morphospecies any instances of sympatric speciation). For "punctuated anagenesis" to occur, the best examples are within the protistan fossil record, especially amongst the radiolarians and foraminifera.
Sorry, John. As a current systematist, I also respectfully beg to differ. We have no way of knowing if those were branching events. If we find two disparate morhpologies in the same locality, we can often suppose that they are separate species (with caveats for sexual and other dimorphism, ecotypes, preservational artifacts), but the appearance of two such morphotypes can often be explained by migration. Or, if geographic sampling is good, as with the Eldredge Phacops instance, we can't tell if there were already two morphologically similar species before one of them experienced punctuated anagenesis. And cryptic/sibling species are very common, and would be even more common if we had only the limited information for extant species that can be gleaned from fossils. I'm certainly not the first to point this out.
John, I did allude to your concerns to an extant in my prior post. The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record. We can quibble as to whether these are in fact clusters of sympatric species, etc. but the patterns are real and can not be easily dismissible as you contend.

Jim Thomerson · 24 January 2011

Most of my taxonomic/systematic work has been on Rivulid killifishes. I don't know of any fossil forms. Something to be said for that.

John Harshman · 24 January 2011

John, I did allude to your concerns to an extant in my prior post. The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record. We can quibble as to whether these are in fact clusters of sympatric species, etc. but the patterns are real and can not be easily dismissible as you contend.
Stasis only means that morphological change is punctuated. (As an aside, I don't think stasis is as well documented as you think, but never mind that part.) At the moment I'm not questioning the pattern of stasis and punctuation. I'm questioning whether punctuation is coincident with speciation. You haven't addressed my point at all so far. By the way, sympatric speciation isn't necessary here. All that's required is that the species be in sympatry when one of them experiences morphological punctuation.

John Kwok · 24 January 2011

John Harshman said:
John, I did allude to your concerns to an extant in my prior post. The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record. We can quibble as to whether these are in fact clusters of sympatric species, etc. but the patterns are real and can not be easily dismissible as you contend.
Stasis only means that morphological change is punctuated. (As an aside, I don't think stasis is as well documented as you think, but never mind that part.) At the moment I'm not questioning the pattern of stasis and punctuation. I'm questioning whether punctuation is coincident with speciation. You haven't addressed my point at all so far. By the way, sympatric speciation isn't necessary here. All that's required is that the species be in sympatry when one of them experiences morphological punctuation.
When I speak of punctuated equilibrium, I am not referring at all to phyletic gradualism/punctuated anagenesis (I am actually skeptical of the latter term, since I am aware of substantial instances of PG within the protistan fossil record.). I am referring to some kind of rapid morphological divergence from an ancestral to a descendant population that would indicate speciation as defined on the basis of morphology.

John Harshman · 25 January 2011

John Kwok said: When I speak of punctuated equilibrium, I am not referring at all to phyletic gradualism/punctuated anagenesis (I am actually skeptical of the latter term, since I am aware of substantial instances of PG within the protistan fossil record.). I am referring to some kind of rapid morphological divergence from an ancestral to a descendant population that would indicate speciation as defined on the basis of morphology.
Ah, so in fact PE is correct by definition, if we use your definition. PE is a theory that most morphological change happens during speciation events. If we define "speciation event" as "episode of morphological change", then PE reduces to a claim that there are episodes of rapid morphological change. That's hardly what Eldredge and Gould had in mind, but it has the advantage of being tautological, and therefore always true. Why you would be skeptical about punctuated anagenesis while embracing punctuated equilibria is beyond me, since under your definition they're the same thing.

SAwells · 25 January 2011

It isn't tautological to theorise that there are episodes of rapid morphological change; whether or not there are such episodes (it looks like there are) is an empirical question.

M.W. · 25 January 2011

MW, since you like the Gospel according to John, how about John 6:46-56? Do you take that literally?

The Pope does, and Benedict XVI, John Paul II, and Pius XII have all said there are no problems with the biological theory of evolution, specifically with regard to the evolution of the human body.

fusilier
James 2:24

Do I take John 6 vs 46-56 literally, the thing about that is if Jesus said it then he meant it, you have to take into consideration why Jesus came and if that was the persuasion of some at that time he would do everything to save a lost soul, but for that way of salvation it is now 2011 years long gone. I take note of what Paul said in 1 Corinthians ch 8 vs 13. At the last supper Jesus used bread and wine for us to remember him by, but first for a person to search their conscience as to their own partaking of the ceremony. If you are hinting that it was through the partaking of that procedure that we became the same human species as he was, then if we eat lamb for example the process would be the same, but I don't think there is any evidence of that, but he was special, he was a healer, so you never know it could be the case, similar to if you were to make a vaccine from some one who had recovered from an illness. Medical processes have developed greatly over the millenniums.

harold · 25 January 2011

John Kwok -
The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record.
Part of the problem is that morphological stasis is at least partly a subjective perception. We don't have good units for measuring it. We probably could make fairly objective methods of measuring genetic change over time within given lineages or subsets of local biospheres, using sampling, with the caveat that the selection of what and who to measure would be a subject of deserved controversy. In my view, the "battle" between PE and "gradualism" is a battle between exaggeratedly extreme straw man versions of each. It's highly reasonable to think that metazoans sometimes radiate rapidly into new niches, and also highly reasonable to think that in some stable environments, highly adapted populations are morphologically stable over long periods of time.

John Harshman · 25 January 2011

SAwells said: It isn't tautological to theorise that there are episodes of rapid morphological change; whether or not there are such episodes (it looks like there are) is an empirical question.
No. But it's tautological to theorize that episodes of rapid morphological change are coincident with speciation, and then define speciation as "an episode of rapid morphological change".

Sonya Troff · 25 January 2011

This comment has been moved to The Bathroom Wall.

John Harshman · 25 January 2011

This comment has been moved to The Bathroom Wall.

John Vanko · 25 January 2011

CMI has posted their article on the Denisovans today:
http://creation.com/denisovan

Pretty much what you'd expect: (paraphrasing in one sentence) "The Denisovans are what we expect from the dispersal after the Flood, proving the Truth of the Bible, but an embarrassment for evilutionists who can't explain them."

harold · 25 January 2011

mrg -

Interestingly, I am familiar with the television version of Terry Pratchett's Hogfather. http://en.wikipedia.org/wiki/Terry_Pratchett's_Hogfather

I happened to come across it accidentally on television. It must have been shown in the US.

Despite my massively above average knowledge of classic British fantasy by US/Canadian, and probably even by UK standards, I had not heard of the books. I had no idea what I was watching but it was quite entertaining.

mrg · 25 January 2011

harold said: I happened to come across it accidentally on television. It must have been shown in the US.
It might have been shown on Sy-Fy Channel, but I picked it up on DVD. I was quite taken with Michelle Dockery as Susan Death ... now she's a high-class Britchick. Surprisingly, she hasn't hit it off in Hollywood yet.

John Kwok · 26 January 2011

John Harshman said:
John Kwok said: When I speak of punctuated equilibrium, I am not referring at all to phyletic gradualism/punctuated anagenesis (I am actually skeptical of the latter term, since I am aware of substantial instances of PG within the protistan fossil record.). I am referring to some kind of rapid morphological divergence from an ancestral to a descendant population that would indicate speciation as defined on the basis of morphology.
Ah, so in fact PE is correct by definition, if we use your definition. PE is a theory that most morphological change happens during speciation events. If we define "speciation event" as "episode of morphological change", then PE reduces to a claim that there are episodes of rapid morphological change. That's hardly what Eldredge and Gould had in mind, but it has the advantage of being tautological, and therefore always true. Why you would be skeptical about punctuated anagenesis while embracing punctuated equilibria is beyond me, since under your definition they're the same thing.
Sorry John, think you need to read again Eldredge and Gould's work. In the very first paper on Punctuated Equilibrium (though the term itself wasn't coined until 1972 by Steve Gould), Eldredge, in a paper published in either 1970 or 1971 proposed that what he saw in his trilobites was some form of allopatric speciation. This is what I think both he and Gould had in mind when they wrote their classic 1972 paper in which they compared and contrasted punctuated equilibrium with phyletic gradualism. Where I am remiss in my earlier comment is that I forgot to mention the analogy from allopatric speciation (Not surprisingly Ernst Mayr would later claim that all punctuated equilibrium is is allopatric speciation as applied to the metazoan fossil record, and to a certain extent he was right.). As for "punctuated anagenesis" there is an extensive body of literature that supports it for Protista (and, if I'm not mistaken, Diatoms too), but more in the sense of true phyletic gradualism over time.

John Kwok · 26 January 2011

harold said: John Kwok -
The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record.
Part of the problem is that morphological stasis is at least partly a subjective perception. We don't have good units for measuring it. We probably could make fairly objective methods of measuring genetic change over time within given lineages or subsets of local biospheres, using sampling, with the caveat that the selection of what and who to measure would be a subject of deserved controversy. In my view, the "battle" between PE and "gradualism" is a battle between exaggeratedly extreme straw man versions of each. It's highly reasonable to think that metazoans sometimes radiate rapidly into new niches, and also highly reasonable to think that in some stable environments, highly adapted populations are morphologically stable over long periods of time.
Neither Eldredge or Gould (or any of their supporters in paleobiology BTW) would claim that you don't see any morphological change within a lineage. Rather, what they have contended is that the net statistical change is zero - hence morphological stasis - and thata any observed variation is really ecophenotypic variation, not the punctuated anagenesis that John Harshman has suggested.

John Kwok · 26 January 2011

Actually, I need to correct myself here. By net zero statistical morphological change, I am not referring to the lineage as a whole, but of the morphospecies under investigation. According to Punctuated Equilibrium then, you would see a rapid divergence from an ancestral population in some kind of branching event - which would be viewed as a speciation event analogous to allopatric speciation - with subsequent morphological change reflective only of ecophenotypic variation as the morphospecies was tracking changing environmental conditions until the time it became extinct:
John Kwok said:
harold said: John Kwok -
The phenomenom of long-term morphological stasis, at least amongst metazoans, is now fairly well established in the fossil record.
Part of the problem is that morphological stasis is at least partly a subjective perception. We don't have good units for measuring it. We probably could make fairly objective methods of measuring genetic change over time within given lineages or subsets of local biospheres, using sampling, with the caveat that the selection of what and who to measure would be a subject of deserved controversy. In my view, the "battle" between PE and "gradualism" is a battle between exaggeratedly extreme straw man versions of each. It's highly reasonable to think that metazoans sometimes radiate rapidly into new niches, and also highly reasonable to think that in some stable environments, highly adapted populations are morphologically stable over long periods of time.
Neither Eldredge or Gould (or any of their supporters in paleobiology BTW) would claim that you don't see any morphological change within a lineage. Rather, what they have contended is that the net statistical change is zero - hence morphological stasis - and thata any observed variation is really ecophenotypic variation, not the punctuated anagenesis that John Harshman has suggested.

Jimmy · 26 January 2011

I've heard quite a lot about Gould's work, but i would really like to read it for myself.

Anybody have a link or perhaps a know a good book i could check out?

mrg · 26 January 2011

Jimmy said: Anybody have a link or perhaps a know a good book i could check out?
A good number of Gould's essays are here: http://www.stephenjaygould.org/library.html I have the oddest suspicion that some will recommend THE STRUCTURE OF EVOLUTIONARY THEORY. "Light reading it is not."

John Harshman · 26 January 2011

John Kwok said: Sorry John, think you need to read again Eldredge and Gould's work. In the very first paper on Punctuated Equilibrium (though the term itself wasn't coined until 1972 by Steve Gould), Eldredge, in a paper published in either 1970 or 1971 proposed that what he saw in his trilobites was some form of allopatric speciation. This is what I think both he and Gould had in mind when they wrote their classic 1972 paper in which they compared and contrasted punctuated equilibrium with phyletic gradualism.
Of course they were. They were in fact proposing a paleontolgoical interpretation of Mayr's peripatric speciation. So? How is that relevant?
Where I am remiss in my earlier comment is that I forgot to mention the analogy from allopatric speciation
It isn't an analogy. It *is* allopatric speciation. But none of this is relevant to our argument. You can't recognize speciation in the fossil record.
As for "punctuated anagenesis" there is an extensive body of literature that supports it for Protista (and, if I'm not mistaken, Diatoms too), but more in the sense of true phyletic gradualism over time.
Punctuated anagenesis isn't phyletic gradualism. It's punctuated. That's why the word "punctuated" is in the name. The question is how you tell PA from PE, i.e. how you tell whether a given morphological change was accompanied simultaneously by speciation. I contend that you can't. You can occasionally tell that what looked rather like a single population has become two morphologically divergent populations. But that could also be a PA event in one (or both) of two formerly sibling species. How can you distinguish sort of event (PE) from the other (prior speciation followed at some indeterminate interval by PA)? There is of course a good model for the latter: character displacement.

Flint · 26 January 2011

how you tell whether a given morphological change was accompanied simultaneously by speciation. I contend that you can’t.

OK, I'm confused. I've seen many different approaches to categorizing speciation, which are not mutually antagonistic, but which are instead useful for different purposes. And one of those purposes has to do with morphological change. Generally, at least as I understand it, morphological diversion in practice requires breeding isolation for some period of time between populations. Or are we saying here that a single breeding population might undergo significant morphological change without speciation, breaking the statis? And this strikes me as problematical - if we have a continuously interbreeding population which undergoes some morphological changes over time, is there EVER a point where we'd say the population "deserves to be called" a new species? I'm starting to wonder whether we have more terms than we have conditions to describe with them.

John Kwok · 26 January 2011

John Harshman said:
John Kwok said: Sorry John, think you need to read again Eldredge and Gould's work. In the very first paper on Punctuated Equilibrium (though the term itself wasn't coined until 1972 by Steve Gould), Eldredge, in a paper published in either 1970 or 1971 proposed that what he saw in his trilobites was some form of allopatric speciation. This is what I think both he and Gould had in mind when they wrote their classic 1972 paper in which they compared and contrasted punctuated equilibrium with phyletic gradualism.
Of course they were. They were in fact proposing a paleontolgoical interpretation of Mayr's peripatric speciation. So? How is that relevant?
Where I am remiss in my earlier comment is that I forgot to mention the analogy from allopatric speciation
It isn't an analogy. It *is* allopatric speciation. But none of this is relevant to our argument. You can't recognize speciation in the fossil record.
As for "punctuated anagenesis" there is an extensive body of literature that supports it for Protista (and, if I'm not mistaken, Diatoms too), but more in the sense of true phyletic gradualism over time.
Punctuated anagenesis isn't phyletic gradualism. It's punctuated. That's why the word "punctuated" is in the name. The question is how you tell PA from PE, i.e. how you tell whether a given morphological change was accompanied simultaneously by speciation. I contend that you can't. You can occasionally tell that what looked rather like a single population has become two morphologically divergent populations. But that could also be a PA event in one (or both) of two formerly sibling species. How can you distinguish sort of event (PE) from the other (prior speciation followed at some indeterminate interval by PA)? There is of course a good model for the latter: character displacement.
John, you are persisting in believing that there are branching events that are more anagenesis than in actual splitting of lineages (which could be viewed as speciation if there is continuous sedimentation going on). If you have finely recorded microstratigraphic sections and see one morphological lineage diverging into two, then that's speciation. If would be punctuated anagenesis if and only if there was one lineage persisting in the stratigraphic section in question and that there were enough net zero statistical change in its morphology that would rule out that lineage tracking ecophenotypic variation.

John Kwok · 26 January 2011

Flint said:

how you tell whether a given morphological change was accompanied simultaneously by speciation. I contend that you can’t.

OK, I'm confused. I've seen many different approaches to categorizing speciation, which are not mutually antagonistic, but which are instead useful for different purposes. And one of those purposes has to do with morphological change. Generally, at least as I understand it, morphological diversion in practice requires breeding isolation for some period of time between populations. Or are we saying here that a single breeding population might undergo significant morphological change without speciation, breaking the statis? And this strikes me as problematical - if we have a continuously interbreeding population which undergoes some morphological changes over time, is there EVER a point where we'd say the population "deserves to be called" a new species? I'm starting to wonder whether we have more terms than we have conditions to describe with them.
Flint what you described in the first paragraph where you see one morphospecies splitting off from another would be viewed as a paleobiological speciation event. As for your second paragraph that describes either punctuated anagenesis or phyletic gradualism. What I find more interesting is long-term persistence of morphological stasis within one or more morphospecies (again with stasis defined as net zero statistical variation not counting for ecophenotypic variation.).

harold · 27 January 2011

John Kwok said -
Neither Eldredge or Gould (or any of their supporters in paleobiology BTW) would claim that you don’t see any morphological change within a lineage. Rather, what they have contended is that the net statistical change is zero - hence morphological stasis - and thata any observed variation is really ecophenotypic variation
I have no reason to disagree with this part of John Kwok's statement. My point is that while it is all very good to talk about amount of morphologic variation, except in cases where there is a simple size change in a defined anatomic element of a lineage, we lack a strong, consistent methodology of objectively quantify morphologic variation. I stand by that point. Stephen Jay Gould is one of my all time favorite writers. I have enjoyed many of his works on a variety of topics. As I noted above, it is my view that it is obvious that, to the human eye, lineages may seem to be "stable" or to "change rapidly" depending on the circumstances, what aspect we have chosen to look for changes in, and how we define "rapidly" and "slowly".

John Harshman · 27 January 2011

John Kwok said: John, you are persisting in believing that there are branching events that are more anagenesis than in actual splitting of lineages (which could be viewed as speciation if there is continuous sedimentation going on). </blockquote. I persist in believing nothing of the sort. I'm claiming nothing more than that the branching events and the morphological divergence are not the same thing, or at least we have no way of telling if morphological divergence is coincident with branching. We can't directly observe splitting of lineages in the fossil record. We can only observe divergence of morphology. That might be happening during or at any time after a branching event, and we have no real way of knowing.
If you have finely recorded microstratigraphic sections and see one morphological lineage diverging into two, then that's speciation.
If you want to define speciation as morphological divergence, yes. But if you are talking about biological species, which is indeed what PE is an attempt to describe, then no. "One morphological lineage" could easily conceal multiple biological species. And if one of them experiences a change, all that does is make the prior separation visible to paleontologists.
If would be punctuated anagenesis if and only if there was one lineage persisting in the stratigraphic section in question and that there were enough net zero statistical change in its morphology that would rule out that lineage tracking ecophenotypic variation.
The second part was quite confused. Punctuated anagenesis would require other than net zero change in morphology. But your conclusion of anagenesis in this case is reasonable. We might suspect PE only if we see two morphotypes where we had previously seen only one. If we see one replaced by another, PE is ruled out. (Of course, the problem of geographic sampling makes even this conclusion ambiguous -- absent adequate sampling, we could be seeing only migration and extinction, with no necessary evolution at all.) And even if we see two morphotypes, we can't say that we have seen branching, since we may have seen only the morphological divergence of prior sibling species.

Jim Thomerson · 27 January 2011

I've not done any of this kind of work myself, but colleagues have shown me work which involved establishing landmarks on an organism, or structure, and seeing how the landmarks positions change in relation to each other with growth, or across a group of species, or whatever. So I have the impression there are accepted methods to quantify morphology.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: John, you are persisting in believing that there are branching events that are more anagenesis than in actual splitting of lineages (which could be viewed as speciation if there is continuous sedimentation going on). </blockquote. I persist in believing nothing of the sort. I'm claiming nothing more than that the branching events and the morphological divergence are not the same thing, or at least we have no way of telling if morphological divergence is coincident with branching. We can't directly observe splitting of lineages in the fossil record. We can only observe divergence of morphology. That might be happening during or at any time after a branching event, and we have no real way of knowing.
If you have finely recorded microstratigraphic sections and see one morphological lineage diverging into two, then that's speciation.
If you want to define speciation as morphological divergence, yes. But if you are talking about biological species, which is indeed what PE is an attempt to describe, then no. "One morphological lineage" could easily conceal multiple biological species. And if one of them experiences a change, all that does is make the prior separation visible to paleontologists.
If would be punctuated anagenesis if and only if there was one lineage persisting in the stratigraphic section in question and that there were enough net zero statistical change in its morphology that would rule out that lineage tracking ecophenotypic variation.
The second part was quite confused. Punctuated anagenesis would require other than net zero change in morphology. But your conclusion of anagenesis in this case is reasonable. We might suspect PE only if we see two morphotypes where we had previously seen only one. If we see one replaced by another, PE is ruled out. (Of course, the problem of geographic sampling makes even this conclusion ambiguous -- absent adequate sampling, we could be seeing only migration and extinction, with no necessary evolution at all.) And even if we see two morphotypes, we can't say that we have seen branching, since we may have seen only the morphological divergence of prior sibling species.
John we are arguing past each other. Didn't you read where I said a paleobiological speciation event? I know full well that Punctuated Equilibrium doesn't account for a complex of sympatic species that may exist due to some kind of niche partitioning or benavior (e. g. one species is diurnal and the other nocturnal). Nor would it account for ring species. Again, what I find most intriguing about this is that there is indeed long-term morphological stasis within a morphospecies that continues from shortly after the "birth" of the species until it goes extinct.

John Kwok · 27 January 2011

Jim Thomerson said: I've not done any of this kind of work myself, but colleagues have shown me work which involved establishing landmarks on an organism, or structure, and seeing how the landmarks positions change in relation to each other with growth, or across a group of species, or whatever. So I have the impression there are accepted methods to quantify morphology.
There's been a rich literature on this, from the likes of Fred Bookstein, the late Les Marcus, and many others.

John Kwok · 27 January 2011

harold said: John Kwok said -
Neither Eldredge or Gould (or any of their supporters in paleobiology BTW) would claim that you don’t see any morphological change within a lineage. Rather, what they have contended is that the net statistical change is zero - hence morphological stasis - and thata any observed variation is really ecophenotypic variation
I have no reason to disagree with this part of John Kwok's statement. My point is that while it is all very good to talk about amount of morphologic variation, except in cases where there is a simple size change in a defined anatomic element of a lineage, we lack a strong, consistent methodology of objectively quantify morphologic variation. I stand by that point. Stephen Jay Gould is one of my all time favorite writers. I have enjoyed many of his works on a variety of topics. As I noted above, it is my view that it is obvious that, to the human eye, lineages may seem to be "stable" or to "change rapidly" depending on the circumstances, what aspect we have chosen to look for changes in, and how we define "rapidly" and "slowly".
See my reply to Jim Thomerson. There does exist such a literature for "objectively qualifying morphological variation that has existed from the early 1980s if not before. The advent of "fast" personal computers and statistical languages/packages like R, S, SPSS and SAS have made this possible.

John Kwok · 27 January 2011

harold said: As I noted above, it is my view that it is obvious that, to the human eye, lineages may seem to be "stable" or to "change rapidly" depending on the circumstances, what aspect we have chosen to look for changes in, and how we define "rapidly" and "slowly".
Stephen Jay Gould was an early pioneer of morphometrics, which was developed subsequently further by a lot of people, of which among those prominent included the likes of Fred Bookstein and Les Marcus. We have an extensive literature that stretches as far back as the early to mid 1960s if not before. However, as I have noted, the advent of "fast" personal computers and the programming languages and statistical packages I have cited (I omitted BMDP and Matlab as those that may still be in use.), have made this more a trivial exercise than many unfamiliar with this field might recognize.

John Harshman · 27 January 2011

John Kwok said: John we are arguing past each other.
I would put it differently. I would say that you don't understand what I'm saying.
Didn't you read where I said a paleobiological speciation event? I know full well that Punctuated Equilibrium doesn't account for a complex of sympatic species that may exist due to some kind of niche partitioning or benavior (e. g. one species is diurnal and the other nocturnal). Nor would it account for ring species. Again, what I find most intriguing about this is that there is indeed long-term morphological stasis within a morphospecies that continues from shortly after the "birth" of the species until it goes extinct.
But if PE refers only to morphospecies, not biological species, it's nearly tautological. Yes, morphospecies experience stasis. If they didn't, they'd be divided up into multiple morphospecies. Stripped of a connection to biological species, PE is reduced to a statement that rates of evolution vary through the history of a lineage. Interesting, but much less interesting than the original claims of PE.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: John we are arguing past each other.
I would put it differently. I would say that you don't understand what I'm saying.
Didn't you read where I said a paleobiological speciation event? I know full well that Punctuated Equilibrium doesn't account for a complex of sympatic species that may exist due to some kind of niche partitioning or benavior (e. g. one species is diurnal and the other nocturnal). Nor would it account for ring species. Again, what I find most intriguing about this is that there is indeed long-term morphological stasis within a morphospecies that continues from shortly after the "birth" of the species until it goes extinct.
But if PE refers only to morphospecies, not biological species, it's nearly tautological. Yes, morphospecies experience stasis. If they didn't, they'd be divided up into multiple morphospecies. Stripped of a connection to biological species, PE is reduced to a statement that rates of evolution vary through the history of a lineage. Interesting, but much less interesting than the original claims of PE.
I strongly disagree. I have referred to paleobiological speciation events, which are true lineage splitting events, (not some persisting lineage in which there is rapid morphological change via some form of punctuated anagenesis or phyletic gradualism), in which one descendant population emerges from an ancestral, much larger, population (as woudl be the case if the descendant population in question is a peripheral isolate of the former, while both continue to persist in the fossil record.). I also agree with you that puncutuated equilibrium does not account for closely related species that are morphologically the same but differ due to some kind of habitat differentiation (e. g. are nocturnal and diurnal for example) or ring species. In fact that is why I have emphasized the term morphospecies. The best examples of punctuated anagenesis or phyletic gradualism can be seen in the marine Protista or Diatom fossil record. In stark contrast, at least for marine metazoans, we see paleobiological speciation events followed by long-term morphological stasis.

John Harshman · 27 January 2011

John Kwok said: I strongly disagree. I have referred to paleobiological speciation events, which are true lineage splitting events, (not some persisting lineage in which there is rapid morphological change via some form of punctuated anagenesis or phyletic gradualism), in which one descendant population emerges from an ancestral, much larger, population (as woudl be the case if the descendant population in question is a peripheral isolate of the former, while both continue to persist in the fossil record.).
How do you know that you're looking at a splitting event, not the subsequent divergence of two species that actually split a while ago?
I also agree with you that puncutuated equilibrium does not account for closely related species that are morphologically the same but differ due to some kind of habitat differentiation (e. g. are nocturnal and diurnal for example) or ring species. In fact that is why I have emphasized the term morphospecies.
Why couldn't punctuated equilibrium apply to sibling species? What I'm saying is that you just can't tell if a morphological change in the fossil record is or is not happening at the same time as a split of lineages. This is a simple claim, but you keep bringing up tangential arguments that have no bearing on that claim. By the way, sibling species may not differ in habitat. If they're sympatric, they must partition resources in some way, but it could be as simple as feeding in different parts of the same tree. And of course there are plenty of allopatric sibling species too.
The best examples of punctuated anagenesis or phyletic gradualism can be seen in the marine Protista or Diatom fossil record. In stark contrast, at least for marine metazoans, we see paleobiological speciation events followed by long-term morphological stasis.
You keep equating punctuated anagenesis with phyletic gradualism. Please stop. And do you consider that a paleobiological speciation event is something quite distinct from a modern speciation event? If so, what makes it different? If you're giving up all claims that a morphological species is also a biological species, you are also giving up claims to PE's main thesis, as well as claims to be able to detect actual splitting events in the fossil record. At most you can determine that a splitting event happened some time prior to observed differentiation. Shall I mention that I also doubt that small, peripherally isolated populations are generally very important in evolution? Mayr's model of speciation isn't a very useful model.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: I strongly disagree. I have referred to paleobiological speciation events, which are true lineage splitting events, (not some persisting lineage in which there is rapid morphological change via some form of punctuated anagenesis or phyletic gradualism), in which one descendant population emerges from an ancestral, much larger, population (as woudl be the case if the descendant population in question is a peripheral isolate of the former, while both continue to persist in the fossil record.).
How do you know that you're looking at a splitting event, not the subsequent divergence of two species that actually split a while ago?
I also agree with you that puncutuated equilibrium does not account for closely related species that are morphologically the same but differ due to some kind of habitat differentiation (e. g. are nocturnal and diurnal for example) or ring species. In fact that is why I have emphasized the term morphospecies.
Why couldn't punctuated equilibrium apply to sibling species? What I'm saying is that you just can't tell if a morphological change in the fossil record is or is not happening at the same time as a split of lineages. This is a simple claim, but you keep bringing up tangential arguments that have no bearing on that claim. By the way, sibling species may not differ in habitat. If they're sympatric, they must partition resources in some way, but it could be as simple as feeding in different parts of the same tree. And of course there are plenty of allopatric sibling species too.
The best examples of punctuated anagenesis or phyletic gradualism can be seen in the marine Protista or Diatom fossil record. In stark contrast, at least for marine metazoans, we see paleobiological speciation events followed by long-term morphological stasis.
You keep equating punctuated anagenesis with phyletic gradualism. Please stop. And do you consider that a paleobiological speciation event is something quite distinct from a modern speciation event? If so, what makes it different? If you're giving up all claims that a morphological species is also a biological species, you are also giving up claims to PE's main thesis, as well as claims to be able to detect actual splitting events in the fossil record. At most you can determine that a splitting event happened some time prior to observed differentiation. Shall I mention that I also doubt that small, peripherally isolated populations are generally very important in evolution? Mayr's model of speciation isn't a very useful model.
1) Again if we are seeing a lineage splitting event in a stratigraphic section that shows continuous deposition (no sign of subequent removal of the sediment), then it has to be seen as a speciation event (I am cautiously using the term paleobiological speciation event just to avoid issues with dealing with ring species and sympatric species), and no, I agree with Eldredge and Gould that such speciation events do indicate the emergence of species as discrete entities that are born and eventually do die. 2) If there are two sibling species emerging from an ancestral species (which persists in the fossil record), then you can't tell them apart if their morphology is the same, which is what you would expect from sympatric speciation. If there are substantial differences between the two, then of course it is possible to have them evolve via punctuated equilibrium. 3) There are different models of allopatric speciation, and one we haven't discussed at length is what one would call vicariant speciation in which you would see a splitting of an ancestral population into two descendant ones. 4) If you have a Protistan or Diatom lineage that is noted by brief, rapid instances of morphological change followed by long-term stasis, and this pattern repeats itself, then it is fair to refer to it as punctuated anagenesis. If, howver, such change is more gradual, then it should be referred to as phyletic gradualism.

John Kwok · 27 January 2011

Some typos which I have corrected:
John Harshman said:
John Kwok said: I strongly disagree. I have referred to paleobiological speciation events, which are true lineage splitting events, (not some persisting lineage in which there is rapid morphological change via some form of punctuated anagenesis or phyletic gradualism), in which one descendant population emerges from an ancestral, much larger, population (as woudl be the case if the descendant population in question is a peripheral isolate of the former, while both continue to persist in the fossil record.).
How do you know that you're looking at a splitting event, not the subsequent divergence of two species that actually split a while ago?
I also agree with you that puncutuated equilibrium does not account for closely related species that are morphologically the same but differ due to some kind of habitat differentiation (e. g. are nocturnal and diurnal for example) or ring species. In fact that is why I have emphasized the term morphospecies.
Why couldn't punctuated equilibrium apply to sibling species? What I'm saying is that you just can't tell if a morphological change in the fossil record is or is not happening at the same time as a split of lineages. This is a simple claim, but you keep bringing up tangential arguments that have no bearing on that claim. By the way, sibling species may not differ in habitat. If they're sympatric, they must partition resources in some way, but it could be as simple as feeding in different parts of the same tree. And of course there are plenty of allopatric sibling species too.
The best examples of punctuated anagenesis or phyletic gradualism can be seen in the marine Protista or Diatom fossil record. In stark contrast, at least for marine metazoans, we see paleobiological speciation events followed by long-term morphological stasis.
You keep equating punctuated anagenesis with phyletic gradualism. Please stop. And do you consider that a paleobiological speciation event is something quite distinct from a modern speciation event? If so, what makes it different? If you're giving up all claims that a morphological species is also a biological species, you are also giving up claims to PE's main thesis, as well as claims to be able to detect actual splitting events in the fossil record. At most you can determine that a splitting event happened some time prior to observed differentiation. Shall I mention that I also doubt that small, peripherally isolated populations are generally very important in evolution? Mayr's model of speciation isn't a very useful model.
1) Again if we are seeing a lineage splitting event in a stratigraphic section that shows continuous deposition (no sign of subsequent removal of the sediment via erosion or any sign of nondeposition), then it has to be seen as a speciation event (I am cautiously using the term paleobiological speciation event just to avoid issues with dealing with ring species and sympatric species), and yes, I agree with Eldredge and Gould that such speciation events do indicate the emergence of species as discrete entities that are born and eventually do die. 2) If there are two sibling species emerging from an ancestral species (which persists in the fossil record), then you can’t tell them apart if their morphology is the same, which is what you would expect from sympatric speciation. If there are substantial differences between the two, then of course it is possible to have them evolve via punctuated equilibrium. 3) There are different models of allopatric speciation, and one we haven’t discussed at length is what one would call vicariant speciation in which you would see a splitting of a large ancestral population into two smaller descendant ones due to the creation of some kind of barrier. 4) If you have a Protistan or Diatom lineage that is noted by brief, rapid instances of morphological change followed by long-term stasis), then it is fair to refer to it as punctuated anagenesis, especially if it repeats itself. If, howver, such change is more gradual, then it should be referred to as phyletic gradualism. In neither instance do you see any genuine lineage splitting, but instead, morphological change within the same lineage, so it would be fair to group the two togetner as evolutionary patterns distinct from punctuated equilibrium.

John Harshman · 27 January 2011

John Kwok said: 1) Again if we are seeing a lineage splitting event
Let's stop right there. What do you mean "lineage splitting event"? What you actually see, at most, is a unimodal distribution of some characteristics gradually (or rapidly, if you prefer) becoming a bimodal distribution. You interpret that as a splitting event. I'm asking how you can say that, when divergence of two species that were already separate, but had previously been similar morphologically, would show exactly the same pattern.
yes, I agree with Eldredge and Gould that such speciation events do indicate the emergence of species as discrete entities that are born and eventually do die.
I know you do. I'm asking how you can make that assumption.
2) If there are two sibling species emerging from an ancestral species (which persists in the fossil record), then you can’t tell them apart if their morphology is the same, which is what you would expect from sympatric speciation. If there are substantial differences between the two, then of course it is possible to have them evolve via punctuated equilibrium.
That's totally opaque as well as irrelevant to our disagreement. I'm not talking about sympatric speciation. Nor, in fact, are sibling species what we would expect, particularly, from sympatric speciation, at least to any greater degree than we would expect from allopatric speciation.
3) There are different models of allopatric speciation, and one we haven’t discussed at length is what one would call vicariant speciation in which you would see a splitting of a large ancestral population into two smaller descendant ones due to the creation of some kind of barrier.
Please stop trying to educate me about things every evolutionary biologist should know already. Assume I know that stuff until I present evidence to the contrary. Again, this is irrelevant too.
4) If you have a Protistan or Diatom lineage that is noted by brief, rapid instances of morphological change followed by long-term stasis), then it is fair to refer to it as punctuated anagenesis, especially if it repeats itself. If, howver, such change is more gradual, then it should be referred to as phyletic gradualism. In neither instance do you see any genuine lineage splitting, but instead, morphological change within the same lineage, so it would be fair to group the two togetner as evolutionary patterns distinct from punctuated equilibrium.
How do you know? How can you distinguish punctuated equilibrium (involving speciation) from punctuated anagenesis (not involving speciation)? Again, this requires the assumption that you can identify single species in the fossil record, and distinguish one species from two sibling species.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: 1) Again if we are seeing a lineage splitting event
Let's stop right there. What do you mean "lineage splitting event"? What you actually see, at most, is a unimodal distribution of some characteristics gradually (or rapidly, if you prefer) becoming a bimodal distribution. You interpret that as a splitting event. I'm asking how you can say that, when divergence of two species that were already separate, but had previously been similar morphologically, would show exactly the same pattern.
yes, I agree with Eldredge and Gould that such speciation events do indicate the emergence of species as discrete entities that are born and eventually do die.
I know you do. I'm asking how you can make that assumption.
2) If there are two sibling species emerging from an ancestral species (which persists in the fossil record), then you can’t tell them apart if their morphology is the same, which is what you would expect from sympatric speciation. If there are substantial differences between the two, then of course it is possible to have them evolve via punctuated equilibrium.
That's totally opaque as well as irrelevant to our disagreement. I'm not talking about sympatric speciation. Nor, in fact, are sibling species what we would expect, particularly, from sympatric speciation, at least to any greater degree than we would expect from allopatric speciation.
3) There are different models of allopatric speciation, and one we haven’t discussed at length is what one would call vicariant speciation in which you would see a splitting of a large ancestral population into two smaller descendant ones due to the creation of some kind of barrier.
Please stop trying to educate me about things every evolutionary biologist should know already. Assume I know that stuff until I present evidence to the contrary. Again, this is irrelevant too.
4) If you have a Protistan or Diatom lineage that is noted by brief, rapid instances of morphological change followed by long-term stasis), then it is fair to refer to it as punctuated anagenesis, especially if it repeats itself. If, howver, such change is more gradual, then it should be referred to as phyletic gradualism. In neither instance do you see any genuine lineage splitting, but instead, morphological change within the same lineage, so it would be fair to group the two togetner as evolutionary patterns distinct from punctuated equilibrium.
How do you know? How can you distinguish punctuated equilibrium (involving speciation) from punctuated anagenesis (not involving speciation)? Again, this requires the assumption that you can identify single species in the fossil record, and distinguish one species from two sibling species.
I think you need to learn something about biostratigraphy, sedimentology as well as invertebrate paleobiology, John and I have neither the time nor patience to educate you here. 1) Again if you have a stratigraphic sequence that demonstrates a continuous record of sedimentation and see in that, the rapid emergence of one population splitting off from another, then it is a speciation event (though I've used the caveat paleobiological speciation event for the reasons I have stated twice already.) 2)You've been the one picking on me about sibling species so I felt compelled to explain what exactly I am referring to. 3) I wrote that more for the benefit of others reading these comments who are unfamiliar with modes of speciation, and also to remind you that punctuated equilibrium may not be primarily a product of peripatric speciation. 4) Look again at my first comment posted here. If you see only rapid evolutionary transformation within a single lineage followed by long-term stasis, then that's clearly an instance of punctuated anagenesis. 5) Again what I find most intriguing about punctuated equilibrium is that it has formally recognized the persistence of long-term morphological stasis within marine metazoans (at least). That and the realization that species are discrete entities through time, who have "births" and "deaths".

John Harshman · 27 January 2011

John Kwok said: I think you need to learn something about biostratigraphy, sedimentology as well as invertebrate paleobiology, John and I have neither the time nor patience to educate you here. Thank you for your condescension. Are you even reading anything I write? I can see no sign of it, if so. Please. I took paleontology from Dave Raup, Dave Jablonski, and Jim Hopson.
1) Again if you have a stratigraphic sequence that demonstrates a continuous record of sedimentation and see in that, the rapid emergence of one population splitting off from another, then it is a speciation event (though I've used the caveat paleobiological speciation event for the reasons I have stated twice already.)
Yes, you have repeated many claims without seeming to notice my objections, much less respond to them. How do you know there is "one population splitting off from another" rather than "one population becoming recognizably distinct from another"?
2)You've been the one picking on me about sibling species so I felt compelled to explain what exactly I am referring to.
Yes, and you responded with irrelevant comments about sympatric speciation. You have never responded to my actual point.
3) I wrote that more for the benefit of others reading these comments who are unfamiliar with modes of speciation, and also to remind you that punctuated equilibrium may not be primarily a product of peripatric speciation.
I need no reminding. (Though if peripatric speciation isn't the usual mode, the original rationale for PE disappears.)
4) Look again at my first comment posted here. If you see only rapid evolutionary transformation within a single lineage followed by long-term stasis, then that's clearly an instance of punctuated anagenesis.
Yes, and I'm asking how you can tell the difference between 1) transformation in a single lineage, when there are sibling species present, one of which doesn't transform, and 2) splitting of lineages, of which one transforms and the other doesn't.
5) Again what I find most intriguing about punctuated equilibrium is that it has formally recognized the persistence of long-term morphological stasis within marine metazoans (at least). That and the realization that species are discrete entities through time, who have "births" and "deaths".
Stasis may be interesting, but it isn't the central claim of PE, which is about speciation. But yes, stasis is interesting. Species as temporally discrete entities is probably hallucination brought on by identification of speciation with punctuation.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: I think you need to learn something about biostratigraphy, sedimentology as well as invertebrate paleobiology, John and I have neither the time nor patience to educate you here. Thank you for your condescension. Are you even reading anything I write? I can see no sign of it, if so. Please. I took paleontology from Dave Raup, Dave Jablonski, and Jim Hopson.
1) Again if you have a stratigraphic sequence that demonstrates a continuous record of sedimentation and see in that, the rapid emergence of one population splitting off from another, then it is a speciation event (though I've used the caveat paleobiological speciation event for the reasons I have stated twice already.)
Yes, you have repeated many claims without seeming to notice my objections, much less respond to them. How do you know there is "one population splitting off from another" rather than "one population becoming recognizably distinct from another"?
2)You've been the one picking on me about sibling species so I felt compelled to explain what exactly I am referring to.
Yes, and you responded with irrelevant comments about sympatric speciation. You have never responded to my actual point.
3) I wrote that more for the benefit of others reading these comments who are unfamiliar with modes of speciation, and also to remind you that punctuated equilibrium may not be primarily a product of peripatric speciation.
I need no reminding. (Though if peripatric speciation isn't the usual mode, the original rationale for PE disappears.)
4) Look again at my first comment posted here. If you see only rapid evolutionary transformation within a single lineage followed by long-term stasis, then that's clearly an instance of punctuated anagenesis.
Yes, and I'm asking how you can tell the difference between 1) transformation in a single lineage, when there are sibling species present, one of which doesn't transform, and 2) splitting of lineages, of which one transforms and the other doesn't.
5) Again what I find most intriguing about punctuated equilibrium is that it has formally recognized the persistence of long-term morphological stasis within marine metazoans (at least). That and the realization that species are discrete entities through time, who have "births" and "deaths".
Stasis may be interesting, but it isn't the central claim of PE, which is about speciation. But yes, stasis is interesting. Species as temporally discrete entities is probably hallucination brought on by identification of speciation with punctuation.
I think you have been all too condescending yourself, John. By your own admission you haven't taken courses in stratigraphy and sedimentology. Again if you have a continuous record of sedimentation within a stratigraphic sequence and see a lineage splitting event in which a new lineage emerges from another, then I'd call that a speciation event. As for your very last comment, may I have your permission to share it with Niles Eldrege? Am sure he'd find it most fascinating.

John Kwok · 27 January 2011

typo, so am reposting here -
John Harshman said:
John Kwok said: I think you need to learn something about biostratigraphy, sedimentology as well as invertebrate paleobiology, John and I have neither the time nor patience to educate you here. Thank you for your condescension. Are you even reading anything I write? I can see no sign of it, if so. Please. I took paleontology from Dave Raup, Dave Jablonski, and Jim Hopson.
1) Again if you have a stratigraphic sequence that demonstrates a continuous record of sedimentation and see in that, the rapid emergence of one population splitting off from another, then it is a speciation event (though I've used the caveat paleobiological speciation event for the reasons I have stated twice already.)
Yes, you have repeated many claims without seeming to notice my objections, much less respond to them. How do you know there is "one population splitting off from another" rather than "one population becoming recognizably distinct from another"?
2)You've been the one picking on me about sibling species so I felt compelled to explain what exactly I am referring to.
Yes, and you responded with irrelevant comments about sympatric speciation. You have never responded to my actual point.
3) I wrote that more for the benefit of others reading these comments who are unfamiliar with modes of speciation, and also to remind you that punctuated equilibrium may not be primarily a product of peripatric speciation.
I need no reminding. (Though if peripatric speciation isn't the usual mode, the original rationale for PE disappears.)
4) Look again at my first comment posted here. If you see only rapid evolutionary transformation within a single lineage followed by long-term stasis, then that's clearly an instance of punctuated anagenesis.
Yes, and I'm asking how you can tell the difference between 1) transformation in a single lineage, when there are sibling species present, one of which doesn't transform, and 2) splitting of lineages, of which one transforms and the other doesn't.
5) Again what I find most intriguing about punctuated equilibrium is that it has formally recognized the persistence of long-term morphological stasis within marine metazoans (at least). That and the realization that species are discrete entities through time, who have "births" and "deaths".
Stasis may be interesting, but it isn't the central claim of PE, which is about speciation. But yes, stasis is interesting. Species as temporally discrete entities is probably hallucination brought on by identification of speciation with punctuation.
I think you have been all too condescending yourself, John. By your own admission you haven’t taken courses in stratigraphy and sedimentology. Again if you have a continuous record of sedimentation within a stratigraphic sequence and see a lineage splitting event in which a new lineage emerges from another, then I’d call that a speciation event. One of the central claims of Punctuated equilibrium is the persistence of morphological stasis over the course of geological time, which I am sure both David Raup and David Jablonski reminded you of. As for your very last comment, may I have your permission to share it with Niles Eldredge? Am sure he’d find it most fascinating. As for these comments of yours: "Yes, and I’m asking how you can tell the difference between 1) transformation in a single lineage, when there are sibling species present, one of which doesn’t transform, and 2) splitting of lineages, of which one transforms and the other doesn’t." 1) Have yet to read of "sibling species" in the Protista and Diatoms and frankly I doubt that they exist. 2) You're trying to play "gotcha" and I'm not interested. In the situation to described, the species that transforms is the descendant of the ancestral species (which doest not transform, but instead, remains in its prior state of morphological stasis.

John Harshman · 27 January 2011

John Kwok said: I think you have been all too condescending yourself, John. By your own admission you haven't taken courses in stratigraphy and sedimentology. Again if you have a continuous record of sedimentation within a stratigraphic sequence and see a lineage splitting event in which a new lineage emerges from another, then I'd call that a speciation event. As for your very last comment, may I have your permission to share it with Niles Eldrege? Am sure he'd find it most fascinating.
Actually, I never said I hadn't. And in fact I have. Look, this could be an interesting discussion if you would bother just once to address my argument directly. How do you distinguish 1) one lineage splitting from another from 2) one member of sibling species pair undergoing anagenesis? And by posting to a public forum, I have shared my comment with anyone who cares to notice it. Now, if you were going to respond rationally to this argument, as I see it you have one option only. You can claim that sibling (or perhaps "cryptic" would be less confusing for you) species are rare. Obviously, the more common they are the more likely you are to miss the true speciation event, and the rarer they are the fewer you miss. Now you have instead chosen two approaches: 1) simply assert that you know best by virtue of your superior brain and 2) simply repeat your initial claim over and over without engaging my argument. Try something else.

John Kwok · 27 January 2011

John Harshman said:
John Kwok said: I think you have been all too condescending yourself, John. By your own admission you haven't taken courses in stratigraphy and sedimentology. Again if you have a continuous record of sedimentation within a stratigraphic sequence and see a lineage splitting event in which a new lineage emerges from another, then I'd call that a speciation event. As for your very last comment, may I have your permission to share it with Niles Eldrege? Am sure he'd find it most fascinating.
Actually, I never said I hadn't. And in fact I have. Look, this could be an interesting discussion if you would bother just once to address my argument directly. How do you distinguish 1) one lineage splitting from another from 2) one member of sibling species pair undergoing anagenesis? And by posting to a public forum, I have shared my comment with anyone who cares to notice it. Now, if you were going to respond rationally to this argument, as I see it you have one option only. You can claim that sibling (or perhaps "cryptic" would be less confusing for you) species are rare. Obviously, the more common they are the more likely you are to miss the true speciation event, and the rarer they are the fewer you miss. Now you have instead chosen two approaches: 1) simply assert that you know best by virtue of your superior brain and 2) simply repeat your initial claim over and over without engaging my argument. Try something else.
John I have answered your question again and again with reference to distinguishing between how one lineage could emerge from another (And I personally would prefer calling them species.) IF YOU HAVE a stratigraphic section with a record of continuous sedimentation. You just don't like my answer. As for your other comments I feel like I am replaying again the neontological versus paleontological furor that erupted when the Eldredge and Gould (1972) paper was published (though there was more a furor when the Gould and Eldredge (1977) and the Gould (1980) papers were published. I really have no interest or time to do just that.

John Harshman · 27 January 2011

One of the central claims of Punctuated equilibrium is the persistence of morphological stasis over the course of geological time, which I am sure both David Raup and David Jablonski reminded you of.
Yes, the claim of general stasis is the part of PE that I think can be salvaged, because it can in principal be tested. I can't say I think its prevalance has been very well tested, but it could be.
1) Have yet to read of "sibling species" in the Protista and Diatoms and frankly I doubt that they exist.
Has anyone been looking for them? You probably have to sequence a large sample and look for divergent clades. It wouldn't be easy.
2) You're trying to play "gotcha" and I'm not interested. In the situation to described, the species that transforms is the descendant of the ancestral species (which doest not transform, but instead, remains in its prior state of morphological stasis.
It's an interesting philosophical question to ponder what the "ancestral species" would be. But the point to make is that the ancestral species wasn't there at the time. It was some time in the past, and you didn't notice the true speciation event because it didn't involve significant morphological change. You still have yet to engage my fundamental question. I'm not trying to play gotcha here; I'm trying to have an argument (the full half hour, not just the 5 minutes). But you won't even notice the question.

John Kwok · 27 January 2011

John Harshman said:
One of the central claims of Punctuated equilibrium is the persistence of morphological stasis over the course of geological time, which I am sure both David Raup and David Jablonski reminded you of.
Yes, the claim of general stasis is the part of PE that I think can be salvaged, because it can in principal be tested. I can't say I think its prevalance has been very well tested, but it could be.
1) Have yet to read of "sibling species" in the Protista and Diatoms and frankly I doubt that they exist.
Has anyone been looking for them? You probably have to sequence a large sample and look for divergent clades. It wouldn't be easy.
2) You're trying to play "gotcha" and I'm not interested. In the situation to described, the species that transforms is the descendant of the ancestral species (which doest not transform, but instead, remains in its prior state of morphological stasis.
It's an interesting philosophical question to ponder what the "ancestral species" would be. But the point to make is that the ancestral species wasn't there at the time. It was some time in the past, and you didn't notice the true speciation event because it didn't involve significant morphological change. You still have yet to engage my fundamental question. I'm not trying to play gotcha here; I'm trying to have an argument (the full half hour, not just the 5 minutes). But you won't even notice the question.
I strongly beg to differ, the paleobiological literature does show now numerous examples of morphological stasis. As for "sibling species" in the Protista and Diatoms, you are talking about two groups well studied by both micropaleontologists and biological oceanographers. Am certain that if they did exist, we would have known about them.

John Harshman · 27 January 2011

John Kwok said: I strongly beg to differ, the paleobiological literature does show now numerous examples of morphological stasis. As for "sibling species" in the Protista and Diatoms, you are talking about two groups well studied by both micropaleontologists and biological oceanographers. Am certain that if they did exist, we would have known about them.
Some day it might be nice to discuss your favorite examples of stasis, but that isn't the main issue we were discussing. And are you really using micropaleontologists as your argument against sibling species? The point about sibling species is that paleontologists can't detect them. Now oceanographers might be better. Do you know of any projects in which oceanographers have sequenced large numbers of supposedly identical individuals? I'd have to say that just amplifying the DNA from a single individual would be a challenge.

John Kwok · 28 January 2011

John Harshman said:
John Kwok said: I strongly beg to differ, the paleobiological literature does show now numerous examples of morphological stasis. As for "sibling species" in the Protista and Diatoms, you are talking about two groups well studied by both micropaleontologists and biological oceanographers. Am certain that if they did exist, we would have known about them.
Some day it might be nice to discuss your favorite examples of stasis, but that isn't the main issue we were discussing. And are you really using micropaleontologists as your argument against sibling species? The point about sibling species is that paleontologists can't detect them. Now oceanographers might be better. Do you know of any projects in which oceanographers have sequenced large numbers of supposedly identical individuals? I'd have to say that just amplifying the DNA from a single individual would be a challenge.
Actually I stand corrected since there is some literature demonstrating the existence of sibling species in foraminifera, radiolaria and diatoms (though the latter seems to be very rare). Such distinctions have been made primarily on the basis of morphometric analysis, with increasingly, work involving genetic analysis. Most of this work, however, dates from the early to mid 1990s to the present.

John Kwok · 28 January 2011

John Harshman said: Some day it might be nice to discuss your favorite examples of stasis, but that isn't the main issue we were discussing. And are you really using micropaleontologists as your argument against sibling species? The point about sibling species is that paleontologists can't detect them. Now oceanographers might be better. Do you know of any projects in which oceanographers have sequenced large numbers of supposedly identical individuals? I'd have to say that just amplifying the DNA from a single individual would be a challenge.
Apparently some paleontologists have been able to detect sibling species, at least for microfossils, based on morphometric analysis. As for a "favorite example" of stasis, you should dig up work by University of Cincinnati invertebrate paleobiologist Carlton E. Brett, who in collaboration with Niles Eldredge and Brett's students, have documented what they are think are examples of "coordinated stasis" amongst differen phyla in Middle Devonian ecosystems occurring in what is now the Appalachians. Moreover, there are well known evolutionary biologists like Douglas Futuyma and Massimo Pigliucci who think that morphological stasis is a real phenomenom that needs to be addressed by their fellow neontologists (Futuyma thinks it can be done within the context of the Modern Synthesis, while Pigliucci is advocating an "Expanded Modern Synthesis".

John Harshman · 28 January 2011

Could you be more specific? On Brett's web site, and judging only from titles, I find only this:

Brett, C.E., Ivany, L., Baugh, H.L., and Wall, P, 2008, Coordinated stasis revisited: Taxonomic and ecologic stability in the Devonian of New York: Paleobiology.

Well, it's a start.

And now, since we apparently agree that cryptic species exist, even in diatoms and forams, would you agree that they present an operational problem for detecting splitting events, that is for distinguishing PE from PA?

John Kwok · 28 January 2011

John Harshman said: Could you be more specific? On Brett's web site, and judging only from titles, I find only this: Brett, C.E., Ivany, L., Baugh, H.L., and Wall, P, 2008, Coordinated stasis revisited: Taxonomic and ecologic stability in the Devonian of New York: Paleobiology. Well, it's a start. And now, since we apparently agree that cryptic species exist, even in diatoms and forams, would you agree that they present an operational problem for detecting splitting events, that is for distinguishing PE from PA?
No, I think we are still in disagreement, especially when I noted that we can see splitting events of morphospecies, with a descendant species emerging from an ancestral one (which is true only for punctuated equilibrium, NOT punctuated anagenesis). I think the problem you cite with regards to cryptic species isn't nearly as serious as you contend. If that was really an important issue, then I highly doubt that such prominent biologists as Douglas Futuyma and Massimo Pigliucci would recognize the importance of morphological stasis.

John Kwok · 28 January 2011

John Kwok said:
John Harshman said: Could you be more specific? On Brett's web site, and judging only from titles, I find only this: Brett, C.E., Ivany, L., Baugh, H.L., and Wall, P, 2008, Coordinated stasis revisited: Taxonomic and ecologic stability in the Devonian of New York: Paleobiology. Well, it's a start. And now, since we apparently agree that cryptic species exist, even in diatoms and forams, would you agree that they present an operational problem for detecting splitting events, that is for distinguishing PE from PA?
No, I think we are still in disagreement, especially when I noted that we can see splitting events of morphospecies, with a descendant species emerging from an ancestral one (which is true only for punctuated equilibrium, NOT punctuated anagenesis). I think the problem you cite with regards to cryptic species isn't nearly as serious as you contend. If that was really an important issue, then I highly doubt that such prominent biologists as Douglas Futuyma and Massimo Pigliucci would recognize the importance of morphological stasis.
And John, before you opt to play semantic word games, under Punctuated Equilibrium, both the descendant and ancestral species would have populations that persist, demonstrating morphological stasis.

John Harshman · 28 January 2011

John Kwok said: No, I think we are still in disagreement, especially when I noted that we can see splitting events of morphospecies, with a descendant species emerging from an ancestral one (which is true only for punctuated equilibrium, NOT punctuated anagenesis). I think the problem you cite with regards to cryptic species isn't nearly as serious as you contend. If that was really an important issue, then I highly doubt that such prominent biologists as Douglas Futuyma and Massimo Pigliucci would recognize the importance of morphological stasis.
So basically your argument here amounts to an appeal to authority. Can't be anything to it, because Futuyma and Pigliucci haven't complained. In fact that's wrong. Stasis is the phenomenon they're talking about. That's different from what you're claiming, which is that stasis is broken only during speciation. It's the association of punctuation with speciation that we're arguing about here. Nothing to do with the reality of stasis. I'll say it again. The fossil record is capable of documenting stasis. It is not capable of documenting whether punctuation is associated (chiefly, or wholly) with speciation, because the fossil record isn't good at letting us recognize speciation. And the reason for this is the existence of cryptic species. You claim to see a descendant species emerging from an ancestral one. I claim that you must first rule out the alternative, that there are two species already present, not easily distinguished by fossils and so presenting the same appearance as a single species, one of which undergoes punctuated anagenesis while the other one doesn't. Note that in this case there is still stasis, and there is still punctuation. The fossil record is capable of identifying both of these. What I claim it can't do is determine that the punctuation is coincident with speciation. I am not playing any semantic word games. But your statement "under Punctuated Equilibrium, both the descendant and ancestral species would have populations that persist, demonstrating morphological stasis" is nonsensical. Under PE, the ancestral species would persist and would show stasis, while the descendant species would undergo a punctuation event during speciation, showing non-stasis. While that's a correct description of PE, it does nothing to advance the claim that PE can be distinguished from PA in the presence of sibling species. Again, if we start with two indistinguishable species and one of them undergoes PA, there is no way to tell that apart from PE. In that case, what looks to you like the ancestral species is, initially, two species. What looks like the speciation event is PA in one of the species. And what looks like the ancestor/descendant pair, finally, is really just the same two species you began with; your ancestor is the species that didn't change, and your descendant is the species that did. But no speciation has happened in this scenario. So: how do you tell your scenario from mine? I claim that you can't. You can only, at best, argue that such cryptic species pairs are rare. Would you like to do that?

John Kwok · 28 January 2011

John Harshman said:
John Kwok said: No, I think we are still in disagreement, especially when I noted that we can see splitting events of morphospecies, with a descendant species emerging from an ancestral one (which is true only for punctuated equilibrium, NOT punctuated anagenesis). I think the problem you cite with regards to cryptic species isn't nearly as serious as you contend. If that was really an important issue, then I highly doubt that such prominent biologists as Douglas Futuyma and Massimo Pigliucci would recognize the importance of morphological stasis.
So basically your argument here amounts to an appeal to authority. Can't be anything to it, because Futuyma and Pigliucci haven't complained. In fact that's wrong. Stasis is the phenomenon they're talking about. That's different from what you're claiming, which is that stasis is broken only during speciation. It's the association of punctuation with speciation that we're arguing about here. Nothing to do with the reality of stasis. I'll say it again. The fossil record is capable of documenting stasis. It is not capable of documenting whether punctuation is associated (chiefly, or wholly) with speciation, because the fossil record isn't good at letting us recognize speciation. And the reason for this is the existence of cryptic species. You claim to see a descendant species emerging from an ancestral one. I claim that you must first rule out the alternative, that there are two species already present, not easily distinguished by fossils and so presenting the same appearance as a single species, one of which undergoes punctuated anagenesis while the other one doesn't. Note that in this case there is still stasis, and there is still punctuation. The fossil record is capable of identifying both of these. What I claim it can't do is determine that the punctuation is coincident with speciation. I am not playing any semantic word games. But your statement "under Punctuated Equilibrium, both the descendant and ancestral species would have populations that persist, demonstrating morphological stasis" is nonsensical. Under PE, the ancestral species would persist and would show stasis, while the descendant species would undergo a punctuation event during speciation, showing non-stasis. While that's a correct description of PE, it does nothing to advance the claim that PE can be distinguished from PA in the presence of sibling species. Again, if we start with two indistinguishable species and one of them undergoes PA, there is no way to tell that apart from PE. In that case, what looks to you like the ancestral species is, initially, two species. What looks like the speciation event is PA in one of the species. And what looks like the ancestor/descendant pair, finally, is really just the same two species you began with; your ancestor is the species that didn't change, and your descendant is the species that did. But no speciation has happened in this scenario. So: how do you tell your scenario from mine? I claim that you can't. You can only, at best, argue that such cryptic species pairs are rare. Would you like to do that?
John Harshman said:
John Kwok said: No, I think we are still in disagreement, especially when I noted that we can see splitting events of morphospecies, with a descendant species emerging from an ancestral one (which is true only for punctuated equilibrium, NOT punctuated anagenesis). I think the problem you cite with regards to cryptic species isn't nearly as serious as you contend. If that was really an important issue, then I highly doubt that such prominent biologists as Douglas Futuyma and Massimo Pigliucci would recognize the importance of morphological stasis.
So basically your argument here amounts to an appeal to authority. Can't be anything to it, because Futuyma and Pigliucci haven't complained. In fact that's wrong. Stasis is the phenomenon they're talking about. That's different from what you're claiming, which is that stasis is broken only during speciation. It's the association of punctuation with speciation that we're arguing about here. Nothing to do with the reality of stasis. I'll say it again. The fossil record is capable of documenting stasis. It is not capable of documenting whether punctuation is associated (chiefly, or wholly) with speciation, because the fossil record isn't good at letting us recognize speciation. And the reason for this is the existence of cryptic species. You claim to see a descendant species emerging from an ancestral one. I claim that you must first rule out the alternative, that there are two species already present, not easily distinguished by fossils and so presenting the same appearance as a single species, one of which undergoes punctuated anagenesis while the other one doesn't. Note that in this case there is still stasis, and there is still punctuation. The fossil record is capable of identifying both of these. What I claim it can't do is determine that the punctuation is coincident with speciation. I am not playing any semantic word games. But your statement "under Punctuated Equilibrium, both the descendant and ancestral species would have populations that persist, demonstrating morphological stasis" is nonsensical. Under PE, the ancestral species would persist and would show stasis, while the descendant species would undergo a punctuation event during speciation, showing non-stasis. While that's a correct description of PE, it does nothing to advance the claim that PE can be distinguished from PA in the presence of sibling species. Again, if we start with two indistinguishable species and one of them undergoes PA, there is no way to tell that apart from PE. In that case, what looks to you like the ancestral species is, initially, two species. What looks like the speciation event is PA in one of the species. And what looks like the ancestor/descendant pair, finally, is really just the same two species you began with; your ancestor is the species that didn't change, and your descendant is the species that did. But no speciation has happened in this scenario. So: how do you tell your scenario from mine? I claim that you can't. You can only, at best, argue that such cryptic species pairs are rare. Would you like to do that?
I don't think you really understand Punctuated Equilibrium at all, because BOTH the ancestor and descendant species would exhibit morphological stasis (In the case of the descendant that would be after its morphological divergence from the ancestral species.). I said when the populations persist, that they would show morphological stasis, NOT during the actual lineage-splitting event, which again, under conditions of continuous sedimentation does represent a speciation event. As for making an "argument from authority", I think you ought to plead guilty too, since you thought it necessary to tell me that you studied paleontology with David Raup, David Jablonski and James Hopson. Do you think I really care? I think this discussion has outlived its usefulness and I bid you a good day.

John Harshman · 28 January 2011

John Kwok said: I don't think you really understand Punctuated Equilibrium at all, because BOTH the ancestor and descendant species would exhibit morphological stasis (In the case of the descendant that would be after its morphological divergence from the ancestral species.). I said when the populations persist, that they would show morphological stasis, NOT during the actual lineage-splitting event, which again, under conditions of continuous sedimentation does represent a speciation event.
I understand PE perfectly well. I just didn't understand what you were trying (failing) to say about it. Nor do I understand, even now, what was relevant about your point. We can all agree that an actual lineage-splitting event is a speciation event. The question is whether you can tell that what you're looking at is in fact a lineage-splitting event. You have consistently ignored that question. "Conditions of continuous sedimentation" have nothing to do with it. Even the best butter won't fix your watch.
As for making an "argument from authority", I think you ought to plead guilty too, since you thought it necessary to tell me that you studied paleontology with David Raup, David Jablonski and James Hopson. Do you think I really care?
That wasn't an argument from authority. It was intended to convince you that I am not a complete novice in the field, so you'd stop trying to explain the obvious to me. Didn't work.
I think this discussion has outlived its usefulness and I bid you a good day.
And all without you ever having addressed, or even appearing to have noticed, the point at issue? Let me repeat it, in case it's unclear to anyone who may be reading. If there are such things as cryptic species, can we reliably distinguish punctuated speciation, starring an ancestor and a descendant species, from punctuated anagenesis, starring two initially cryptic species, one of which undergoes punctuated anagenesis? I say we can't. That's what "cryptic" means. Bye.

John Kwok · 28 January 2011

This is my last comment (I hope on this thread), but anyone who reads my discussion with John Harshman should understand that he has failed to understand my references to biostratigraphy, stratigraphy and sedimentology. Had he demonstated some understanding, I strongly doubt that we'd be at such an impasse. As for his insistance on cryptic species, his an argument based partly on incredulity, since most metazoan taxa are not comprised of closely related cryptic sibling species. As I have alluded to a recent comment, the existence of cryptic species in Protists and Diatoms is relatively rare (with Diatoms being the rarest).

John Harshman · 29 January 2011

John Kwok said: This is my last comment (I hope on this thread),
We may all hope, but experience seems to dictate that you must always have the last word.
but anyone who reads my discussion with John Harshman should understand that he has failed to understand my references to biostratigraphy, stratigraphy and sedimentology.
What a pity, then, that you never explained yourself, but merely repeated those words. What exactly didn't I understand? We may never know.
Had he demonstated some understanding, I strongly doubt that we'd be at such an impasse. As for his insistance on cryptic species, his an argument based partly on incredulity, since most metazoan taxa are not comprised of closely related cryptic sibling species. As I have alluded to a recent comment, the existence of cryptic species in Protists and Diatoms is relatively rare (with Diatoms being the rarest).
Finally, an attempt at the real question. Fitting that it should be after you've already declared victory and gone home. In my experience, cryptic species tend to appear when you start looking for them seriously, using the right sort of data. You could of course be right with regard to any particular taxon, but as far as I can see the simpler the apparent morphology, the more a problem cryptic species might be. But thanks for finally responding to the question. That helps. And thanks for the reference you almost provided, to Brett.

John Kwok · 29 January 2011

Sorry John, but for someone who claimed to know Geraat Vermeij well enough to tell if he always gave his lectures from his memory (and also to express your disdain for him simply because he's not enthusiastic about cladistics - I happen to think he's mistaken here myself - but I won't "crucify" him for his indifference toward cladistics, which, I suspect, is what you would prefer to do to Vermeij if it was entirely up to you.), you are once again ignoring the important points with respect to biostratigraphy, stratigraphy and sedimentology that I have been trying to make here, and since you insist on refusing to acknowledging them, then you have no business in not respecting my contention that, under conditions where continuous sedimentation has been preseved in the geological column, then it is possible to identify real speciation events in the fossil record and distinguish them from either instances of the first appearance of immigrant taxa or some form of ecophenotypic variation.

I didn't have a Brett reference handy, but instead, only referred you in more general terms, to the work he has been doing with his students and Niles Eldredge. Maybe after you read that, I might be interested in resuming our discussion, but not until then.

John Kwok · 29 January 2011

John Harshman said: Finally, an attempt at the real question. Fitting that it should be after you've already declared victory and gone home. In my experience, cryptic species tend to appear when you start looking for them seriously, using the right sort of data. You could of course be right with regard to any particular taxon, but as far as I can see the simpler the apparent morphology, the more a problem cryptic species might be. But thanks for finally responding to the question. That helps. And thanks for the reference you almost provided, to Brett.
Again, you've made a mountain out of a molehill with regards to cryptic species. I don't think it is as serious an issue as you might contend.

Jim Thomerson · 29 January 2011

I've examined a lot fish skeletons. There are sister species which have identical skeletons, species which show considerable skeletal variation, and any number of uninformative bones. With fragmented skeletons, it might be very difficult to correctly recognize species. There is a reason paleontologists count genera or families when talking about comparative diversities.

John Harshman · 29 January 2011

John Kwok said: Sorry John,
I'm beginning to suspect you aren't really sorry.
but for someone who claimed to know Geraat Vermeij well enough to tell if he always gave his lectures from his memory (and also to express your disdain for him simply because he's not enthusiastic about cladistics - I happen to think he's mistaken here myself - but I won't "crucify" him for his indifference toward cladistics, which, I suspect, is what you would prefer to do to Vermeij if it was entirely up to you.),
Wow, that was a long, irrelevant digression. And in fact every claim you make in it is false to boot.
you are once again ignoring the important points with respect to biostratigraphy, stratigraphy and sedimentology that I have been trying to make here, and since you insist on refusing to acknowledging them, then you have no business in not respecting my contention that, under conditions where continuous sedimentation has been preseved in the geological column, then it is possible to identify real speciation events in the fossil record and distinguish them from either instances of the first appearance of immigrant taxa or some form of ecophenotypic variation.
If the last part of your sentence is indeed all you're saying about biostratigraphy, stratigraphy, and sedimentology, then I don't agree. Identifying real speciation events under these conditions requires a) ruling out cryptic species, which is impossible, and b) ruling out immigration, which requires geographic sampling good enough to cover all possible sources of immigrants. I'll forget the ecophenotypic variation if you will. But you need to explain why the first two are not a problem, and this requires more than just repeating the mantras of "biostratigraphy, stratigraphy, and sedimentology" and "conditions where continuous sedimentation has been preseved in the geological column" over and over.
I didn't have a Brett reference handy, but instead, only referred you in more general terms, to the work he has been doing with his students and Niles Eldredge. Maybe after you read that, I might be interested in resuming our discussion, but not until then.
Is there only the one paper? I can't find any more, and you implied there was a large body of work. Can you help?

John Harshman · 29 January 2011

John Kwok said: Again, you've made a mountain out of a molehill with regards to cryptic species. I don't think it is as serious an issue as you might contend.
Excellent. You have finally begun an attempt to address my claims. So, how would we know if it's a serious issue? Probably by using modern taxa as a guide. In birds, it's often very hard to tell closely related species apart, even with complete skeletons, which you seldom get in fossils. I once worked in a lab next to a nemertine systematist, who, every time he looked at a new beach, found as many as a dozen new, cryptic species, that he couldn't distinguish at all morphologically. It would be nice to find a review of cryptic species across taxa, but I don't know of any. But based on my personal experience, I consider it a major problem, especially for fossils, for which we have limited data to discriminate species.

John Kwok · 29 January 2011

John,

Carl Brett, Niles Eldredge, and Brett's students (I believe one of Eldredge's too) have published a number of papers on their work, but I don't have the references handy. You'll have to dig it for yourself (Not trying to be obtuse but I'm actually off to hear Massimo Pigliucci speak in short while and then, after that, have other business to do in Manhattan.).

When you've demonstrated some familiarity with stratigraphy and sedimentology, then I'll converse with you more on this.

John Kwok · 29 January 2011

i actually tried to address your points at the very onset where we started sparring, but you were too busy pontificating:
John Harshman said:
John Kwok said: Again, you've made a mountain out of a molehill with regards to cryptic species. I don't think it is as serious an issue as you might contend.
Excellent. You have finally begun an attempt to address my claims. So, how would we know if it's a serious issue? Probably by using modern taxa as a guide. In birds, it's often very hard to tell closely related species apart, even with complete skeletons, which you seldom get in fossils. I once worked in a lab next to a nemertine systematist, who, every time he looked at a new beach, found as many as a dozen new, cryptic species, that he couldn't distinguish at all morphologically. It would be nice to find a review of cryptic species across taxa, but I don't know of any. But based on my personal experience, I consider it a major problem, especially for fossils, for which we have limited data to discriminate species.

John Kwok · 29 January 2011

John Harshman said: But based on my personal experience, I consider it a major problem, especially for fossils, for which we have limited data to discriminate species.
Would you care to explain that to generations of systematic paleobiologists, John? Your comment is utterly without merit and is absurd. Am sure David Raup, David Jablonski and James Hopson would be in agreement with me (I'm merely mentioning them since you were so kind to inform me that you had studied paleontology with them.).

John Harshman · 29 January 2011

John Kwok said:Carl Brett, Niles Eldredge, and Brett's students (I believe one of Eldredge's too) have published a number of papers on their work, but I don't have the references handy.
Brett's online CV only mentions one of them, unless the titles aren't very suggestive. I'm a bit stuck.

John Harshman · 29 January 2011

John Kwok said: Would you care to explain that to generations of systematic paleobiologists, John? Your comment is utterly without merit and is absurd. Am sure David Raup, David Jablonski and James Hopson would be in agreement with me (I'm merely mentioning them since you were so kind to inform me that you had studied paleontology with them.).
So your whole argument here is just name-dropping, while reading minds. Hey, and they're my names you're dropping. I'm hardly the first person to bring up this problem. It's surprising you haven't run into it before. And that isn't an answer.

John Harshman · 29 January 2011

John K:

While I'm here, I would like to clarify one issue and explain another.

First, let's get clear on what we're arguing about. Would you agree that if cryptic species are common, that would make it difficult to distinguish punctuated speciation from punctuated anagenesis using the fossil record?

Second, let me drop another couple of names. I first encountered the cryptic species problem (which I believe I've mentioned I am not making up) in a class on speciation team-taught by Jerry Coyne and Paul Sereno. I wish I could remember whose paper it was; there was a nice figure that explained it simply. But the case is also stated nicely in the book Genetics, Paleontology, and Macroevolution by Jeffrey S. Levinton, which I recommend quite aside from his treatment of this particular point.

John Kwok · 30 January 2011

John Harshman said:
John Kwok said: Would you care to explain that to generations of systematic paleobiologists, John? Your comment is utterly without merit and is absurd. Am sure David Raup, David Jablonski and James Hopson would be in agreement with me (I'm merely mentioning them since you were so kind to inform me that you had studied paleontology with them.).
So your whole argument here is just name-dropping, while reading minds. Hey, and they're my names you're dropping. I'm hardly the first person to bring up this problem. It's surprising you haven't run into it before. And that isn't an answer.
Species have been well defined by generations of paleobiologists John, especially those working in micropaleontology and invertebrate paleobiology. I suggest you start looking at the history of paleontology instead of telling me who taught you paleontology at the University of Chicago. Your point is utterly without merit, unless you are discussing vertebrate paleobiology.

John Kwok · 30 January 2011

John Harshman said: John K: While I'm here, I would like to clarify one issue and explain another. First, let's get clear on what we're arguing about. Would you agree that if cryptic species are common, that would make it difficult to distinguish punctuated speciation from punctuated anagenesis using the fossil record? Second, let me drop another couple of names. I first encountered the cryptic species problem (which I believe I've mentioned I am not making up) in a class on speciation team-taught by Jerry Coyne and Paul Sereno. I wish I could remember whose paper it was; there was a nice figure that explained it simply. But the case is also stated nicely in the book Genetics, Paleontology, and Macroevolution by Jeffrey S. Levinton, which I recommend quite aside from his treatment of this particular point.
I strongly disagree since I have been careful to refer to morphospecies, from the very beginning, in case you haven't noticed. As for Levinton's work - I have ample respect for it - but I think here he's mistaken.

John Kwok · 30 January 2011

John Harshman said:
John Kwok said:Carl Brett, Niles Eldredge, and Brett's students (I believe one of Eldredge's too) have published a number of papers on their work, but I don't have the references handy.
Brett's online CV only mentions one of them, unless the titles aren't very suggestive. I'm a bit stuck.
Am certain Carl has published other work, but don't quote me on this. He and his students have been working on this since the early 1990s.

John Kwok · 30 January 2011

John Harshman said:
John Kwok said:Carl Brett, Niles Eldredge, and Brett's students (I believe one of Eldredge's too) have published a number of papers on their work, but I don't have the references handy.
Brett's online CV only mentions one of them, unless the titles aren't very suggestive. I'm a bit stuck.
I might also add that I have other, more immediate, priorities than compiling a Carl Brett bibliography on your behalf, of which the most pressing one is finishing a revised draft of an unpublished novel that I expect to start shopping to potential literary agents sometime next month. As far as I am concerned, this discussion is now over. Hope yours is a good day, John.

Deklane · 30 January 2011

John Kwok said: I might also add that I have other, more immediate, priorities than compiling a Carl Brett bibliography on your behalf, of which the most pressing one is finishing a revised draft of an unpublished novel that I expect to start shopping to potential literary agents sometime next month. As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
There's a novel here all right. I envision it told from the standpoint of a normal mortal caught in the crossfire between two mighty wizards flinging arcane magical spells at each other along with elegant insults that may sound like professions of deep respect but have hidden zingers. The observer doesn't know what the fight is about or why the wizards are mad at each other, he doesn't understand the spells or even the insults, but the fireworks are certainly impressive to watch.

John Kwok · 30 January 2011

Deklane said:
John Kwok said: I might also add that I have other, more immediate, priorities than compiling a Carl Brett bibliography on your behalf, of which the most pressing one is finishing a revised draft of an unpublished novel that I expect to start shopping to potential literary agents sometime next month. As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
There's a novel here all right. I envision it told from the standpoint of a normal mortal caught in the crossfire between two mighty wizards flinging arcane magical spells at each other along with elegant insults that may sound like professions of deep respect but have hidden zingers. The observer doesn't know what the fight is about or why the wizards are mad at each other, he doesn't understand the spells or even the insults, but the fireworks are certainly impressive to watch.
No, mine is a near future alternative history post-cyberpunk novel set in the USA and Europe. If I wanted to write what you've described, then I would have written it.

Deklane · 30 January 2011

John Kwok said:
Deklane said:
John Kwok said: I might also add that I have other, more immediate, priorities than compiling a Carl Brett bibliography on your behalf, of which the most pressing one is finishing a revised draft of an unpublished novel that I expect to start shopping to potential literary agents sometime next month. As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
There's a novel here all right. I envision it told from the standpoint of a normal mortal caught in the crossfire between two mighty wizards flinging arcane magical spells at each other along with elegant insults that may sound like professions of deep respect but have hidden zingers. The observer doesn't know what the fight is about or why the wizards are mad at each other, he doesn't understand the spells or even the insults, but the fireworks are certainly impressive to watch.
No, mine is a near future alternative history post-cyberpunk novel set in the USA and Europe. If I wanted to write what you've described, then I would have written it.
Er... you did write it. Just now.

John Kwok · 30 January 2011

Deklane said:
John Kwok said:
Deklane said:
John Kwok said: I might also add that I have other, more immediate, priorities than compiling a Carl Brett bibliography on your behalf, of which the most pressing one is finishing a revised draft of an unpublished novel that I expect to start shopping to potential literary agents sometime next month. As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
There's a novel here all right. I envision it told from the standpoint of a normal mortal caught in the crossfire between two mighty wizards flinging arcane magical spells at each other along with elegant insults that may sound like professions of deep respect but have hidden zingers. The observer doesn't know what the fight is about or why the wizards are mad at each other, he doesn't understand the spells or even the insults, but the fireworks are certainly impressive to watch.
No, mine is a near future alternative history post-cyberpunk novel set in the USA and Europe. If I wanted to write what you've described, then I would have written it.
Er... you did write it. Just now.
Mere wishful thinking on your part Deklane. If I want to read such a fantasy, I'll turn to J. K. Rowling and especially, J. R. R. Tolkien. Trust me, I couldn't write anything comparable to theirs.

Deklane · 30 January 2011

When will I learn to be quiet and let the grown-ups talk? I was just expressing in humorous terms my reaction to the exchange of views you and John Harshman just had. Way over my head but fascinating to watch. You two might as well be mighty wizards engaging in combat over matters I know nothing about (Clarke's line about advanced science seeming like magic appropriate here). But when it counts you're on the same side, and I'll leave it at that.

John Harshman · 30 January 2011

John Kwok said: Species have been well defined by generations of paleobiologists John, especially those working in micropaleontology and invertebrate paleobiology. I suggest you start looking at the history of paleontology instead of telling me who taught you paleontology at the University of Chicago. Your point is utterly without merit, unless you are discussing vertebrate paleobiology.
Oddly enough, you also say:
I strongly disagree since I have been careful to refer to morphospecies, from the very beginning, in case you haven’t noticed. As for Levinton’s work - I have ample respect for it - but I think here he’s mistaken.
So when I say it, I'm ignorant, but when Levinton says it, you have great respect. Interesting. Now if you really meant to refer only to morphospecies, we would have no argument. But in that case you couldn't also be talking about lineage splitting, since lineage splitting refers to biological species. A single morphospecies can include several biological species, which is my point, and a new morphospecies can arise without any splitting event, which is also my point.
As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
By my count, this is the third time the discussion has been over. We'll see if third time is the charm.

John Kwok · 30 January 2011

John Harshman said:
John Kwok said: Species have been well defined by generations of paleobiologists John, especially those working in micropaleontology and invertebrate paleobiology. I suggest you start looking at the history of paleontology instead of telling me who taught you paleontology at the University of Chicago. Your point is utterly without merit, unless you are discussing vertebrate paleobiology.
Oddly enough, you also say:
I strongly disagree since I have been careful to refer to morphospecies, from the very beginning, in case you haven’t noticed. As for Levinton’s work - I have ample respect for it - but I think here he’s mistaken.
So when I say it, I'm ignorant, but when Levinton says it, you have great respect. Interesting. Now if you really meant to refer only to morphospecies, we would have no argument. But in that case you couldn't also be talking about lineage splitting, since lineage splitting refers to biological species. A single morphospecies can include several biological species, which is my point, and a new morphospecies can arise without any splitting event, which is also my point.
As far as I am concerned, this discussion is now over. Hope yours is a good day, John.
By my count, this is the third time the discussion has been over. We'll see if third time is the charm.
Levinton is in the distinct minority and he has made much of the same arguments since the mid 1970s. Again, you need to follow this: I reject his criticisms of Punctuated Equilibrium, especially when there major evolutionary biologists, including several of his Stony Brook colleagues, who accept tis reality (though they do not go as far as Eldredge and Gould and Pigliucci might in suggesting that we are in need of an "Extended Modern Synthesis". Again I am sorry, but this discussion really has to end. I have a very bad cold which need to shake off and must return to my writing.

John Harshman · 30 January 2011

John Kwok said: Levinton is in the distinct minority and he has made much of the same arguments since the mid 1970s. Again, you need to follow this: I reject his criticisms of Punctuated Equilibrium, especially when there major evolutionary biologists, including several of his Stony Brook colleagues, who accept tis reality (though they do not go as far as Eldredge and Gould and Pigliucci might in suggesting that we are in need of an "Extended Modern Synthesis". Again I am sorry, but this discussion really has to end. I have a very bad cold which need to shake off and must return to my writing.

John Kwok · 30 January 2011

John I said that morphospecies from the very outset could hide species complexes of cryptic species, sympatrically-derived species and ring species. All you have done is recycle arguments that Levinton and a handful of others, such as the late Thomas J. M. Schopf (a former colleague of Raup's at Chicago), were saying from the early 70s to early 80s. Sorry I have no interest in playing that game, and moreover, if you are so interested in quote mining me, then you have ample time to dig up work by Brett, Eldredge and their students, including one of Carl's former students, an Evangelical Protestant Christian, Keith Miller, who was actively involved in the Kansas Board of Education debates in the prior decade.

John Harshman · 30 January 2011

Whoops.
John Kwok said: Levinton is in the distinct minority and he has made much of the same arguments since the mid 1970s. Again, you need to follow this: I reject his criticisms of Punctuated Equilibrium, especially when there major evolutionary biologists, including several of his Stony Brook colleagues, who accept tis reality (though they do not go as far as Eldredge and Gould and Pigliucci might in suggesting that we are in need of an "Extended Modern Synthesis".
It seems to me that your entire argument here is appeal to authority. You call me ignorant because I'm not famous enough. You respect Levinton, although he says the same I do, because he's famous enough. But you reject him because he disagrees with Eldredge, Gould, Pigliucci, and various Stony Brook biologists, who are apparently even more famous. And you don't mention even one actual argument.
Again I am sorry, but this discussion really has to end. I have a very bad cold which need to shake off and must return to my writing.
Get well soon. But nobody is forcing you to keep this going. All you have to do is stop.

John Harshman · 30 January 2011

John Kwok said: John I said that morphospecies from the very outset could hide species complexes of cryptic species, sympatrically-derived species and ring species. All you have done is recycle arguments that Levinton and a handful of others, such as the late Thomas J. M. Schopf (a former colleague of Raup's at Chicago), were saying from the early 70s to early 80s. Sorry I have no interest in playing that game, and moreover, if you are so interested in quote mining me, then you have ample time to dig up work by Brett, Eldredge and their students, including one of Carl's former students, an Evangelical Protestant Christian, Keith Miller, who was actively involved in the Kansas Board of Education debates in the prior decade.
You display a distinct stripe of paranoia. Nobody is quote-mining you. I've tried to engage your argument honestly, as I see it. But I don't see you doing much. Nor have you answered my claims other than to reject them out of hand, and drop names of people who may agree with you, without attempting to explain why my argument is wrong. Feel free not to reply.

John Kwok · 30 January 2011

Nope it is not an argument based on authority, John. Really advise you learn something about stratigraphy and sedimentology. As for Schopf, he edited and published the 1972 Eldredge and Gould punctuated equilibrium paper for a symposium volume published by W. H. Freeman, and still regarded Gould as a friend when he died unexpectedly from a heart attack in the early 1980s. As for Levinton, his wife has worked closely with Eldredge in the past (So purely for personal reasons I have no interest in attacking him.).

John Kwok · 30 January 2011

John Harshman said:
John Kwok said: John I said that morphospecies from the very outset could hide species complexes of cryptic species, sympatrically-derived species and ring species. All you have done is recycle arguments that Levinton and a handful of others, such as the late Thomas J. M. Schopf (a former colleague of Raup's at Chicago), were saying from the early 70s to early 80s. Sorry I have no interest in playing that game, and moreover, if you are so interested in quote mining me, then you have ample time to dig up work by Brett, Eldredge and their students, including one of Carl's former students, an Evangelical Protestant Christian, Keith Miller, who was actively involved in the Kansas Board of Education debates in the prior decade.
You display a distinct stripe of paranoia. Nobody is quote-mining you. I've tried to engage your argument honestly, as I see it. But I don't see you doing much. Nor have you answered my claims other than to reject them out of hand, and drop names of people who may agree with you, without attempting to explain why my argument is wrong. Feel free not to reply.
Sorry John, I have sought to answer your questions again and again, but you seem disinterested in paying heed to what I have said. As for quote-mining, you have done that - or have at least come close - by cutting and pasting comments I had written.

John Harshman · 31 January 2011

John Kwok said: Nope it is not an argument based on authority, John. Really advise you learn something about stratigraphy and sedimentology.
So, to summarize, I'm an ignorant idiot. Stratigraphy would resolve my question, but I'm so ignorant you can't explain why or how. Levinton, who agrees with me, is not ignorant, but is wrong for unspecified reasons.
As for Schopf, he edited and published the 1972 Eldredge and Gould punctuated equilibrium paper for a symposium volume published by W. H. Freeman, and still regarded Gould as a friend when he died unexpectedly from a heart attack in the early 1980s. As for Levinton, his wife has worked closely with Eldredge in the past (So purely for personal reasons I have no interest in attacking him.).
So, because people are friends or acquaintances, they must either agree or at least not explain why they disagree. Got it.
Sorry John, I have sought to answer your questions again and again, but you seem disinterested in paying heed to what I have said. As for quote-mining, you have done that - or have at least come close - by cutting and pasting comments I had written.
Please stop saying you're sorry when you aren't sorry. "Disinterested" doesn't mean what you think it does. You have in fact not answered my questions. And if you can show where I have quote-mined you -- for which cutting and pasting is not sufficient, the crucial criterion being distortion of meaning -- please do so. Or, if your cold and your novel are too pressing, just stop responding.

Malchus · 31 January 2011

John Harshman is quite correct: you have never offered a specific counter-argument. Continually repeating that some field of study refutes his point is insufficient, and like your continual name-dropping, childish. Refute him, or simply admit that you jumped the gun in error.
John Kwok said:
John Harshman said:
John Kwok said: John I said that morphospecies from the very outset could hide species complexes of cryptic species, sympatrically-derived species and ring species. All you have done is recycle arguments that Levinton and a handful of others, such as the late Thomas J. M. Schopf (a former colleague of Raup's at Chicago), were saying from the early 70s to early 80s. Sorry I have no interest in playing that game, and moreover, if you are so interested in quote mining me, then you have ample time to dig up work by Brett, Eldredge and their students, including one of Carl's former students, an Evangelical Protestant Christian, Keith Miller, who was actively involved in the Kansas Board of Education debates in the prior decade.
You display a distinct stripe of paranoia. Nobody is quote-mining you. I've tried to engage your argument honestly, as I see it. But I don't see you doing much. Nor have you answered my claims other than to reject them out of hand, and drop names of people who may agree with you, without attempting to explain why my argument is wrong. Feel free not to reply.
Sorry John, I have sought to answer your questions again and again, but you seem disinterested in paying heed to what I have said. As for quote-mining, you have done that - or have at least come close - by cutting and pasting comments I had written.

Malchus · 31 January 2011

Kwok, at the moment you are coming across as someone so unwilling to admit error that you are willing to simply obfuscate until the other person gives up in disgust. If you have a specific rebuttal, make it. I don't believe that you can, actually.

John Kwok · 31 January 2011

Malchus said: Kwok, at the moment you are coming across as someone so unwilling to admit error that you are willing to simply obfuscate until the other person gives up in disgust. If you have a specific rebuttal, make it. I don't believe that you can, actually.
I have told John Harshman that there are stratigraphic sections which document continuous sedimentation, and it is in those locations where one could see lineage-splitting events that could be seen as examples of speciation (Assuming that you also sample broadly enough to ensure that you are not picking up the first appearance of some immigrant taxon.). He doesn't have a background in stratigraphy and sedimentology and doesn't like my answer. I also said that since I view these as morphospecies, that it is possible that we are missing out on cases of cryptic species or of subsequently sympatric speciation. However, I disagree with him that the problem is as serious as he contends. Neither you nor John Harshman have a background in stratigraphy and sedimentology and therefore, you aren't qualified to criticize me, or even resort to ad hominem attacks. As far as I am concerned this discussion is now over.

Malchus · 31 January 2011

Kwok, neither of us had engaged in any ad hominems at all - your ignorance of that particular fallacy is interesting.

Malchus · 31 January 2011

And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.

John Kwok · 31 January 2011

Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.

John Harshman · 31 January 2011

John Kwok said: I have told John Harshman that there are stratigraphic sections which document continuous sedimentation, and it is in those locations where one could see lineage-splitting events that could be seen as examples of speciation (Assuming that you also sample broadly enough to ensure that you are not picking up the first appearance of some immigrant taxon.). He doesn't have a background in stratigraphy and sedimentology and doesn't like my answer.
But I do have a background in stratigraphy and sedimentology. At least I took a course in it. The reason I don't like your answer is that it isn't an answer. You would have to explain how to distinguish splitting from anagenesis in one of two or more cryptic species.
I also said that since I view these as morphospecies, that it is possible that we are missing out on cases of cryptic species or of subsequently sympatric speciation.
But if they're morphospecies, they have no necessary connection to splitting events. If there are cryptic species, a new morphospecies, even with a perfect depositional record, could be the result of something other than a splitting event.
However, I disagree with him that the problem is as serious as he contends.
I'm not even sure you understand what the problem is. At least you've been dancing around it for quite a while without a straight answer. Now we could have a real discussion about the crux of it all, i.e. how common cryptic species are. That is in fact the only relevant matter to discuss, but you don't seem to realize that.
Neither you nor John Harshman have a background in stratigraphy and sedimentology and therefore, you aren't qualified to criticize me, or even resort to ad hominem attacks.
Stratigraphy and sedimentology aren't relevant to the central question, even if I didn't know anything about them. And I do anyway. Still, you could try to educate us. What about stratigraphy, etc., renders the cryptic species problem nonexistent?
As far as I am concerned this discussion is now over.
...for the fifth time, by my count, though I may have missed one or two.

John Kwok · 31 January 2011

John Harshman said:
John Kwok said: I have told John Harshman that there are stratigraphic sections which document continuous sedimentation, and it is in those locations where one could see lineage-splitting events that could be seen as examples of speciation (Assuming that you also sample broadly enough to ensure that you are not picking up the first appearance of some immigrant taxon.). He doesn't have a background in stratigraphy and sedimentology and doesn't like my answer.
But I do have a background in stratigraphy and sedimentology. At least I took a course in it. The reason I don't like your answer is that it isn't an answer. You would have to explain how to distinguish splitting from anagenesis in one of two or more cryptic species.
I also said that since I view these as morphospecies, that it is possible that we are missing out on cases of cryptic species or of subsequently sympatric speciation.
But if they're morphospecies, they have no necessary connection to splitting events. If there are cryptic species, a new morphospecies, even with a perfect depositional record, could be the result of something other than a splitting event.
However, I disagree with him that the problem is as serious as he contends.
I'm not even sure you understand what the problem is. At least you've been dancing around it for quite a while without a straight answer. Now we could have a real discussion about the crux of it all, i.e. how common cryptic species are. That is in fact the only relevant matter to discuss, but you don't seem to realize that.
Neither you nor John Harshman have a background in stratigraphy and sedimentology and therefore, you aren't qualified to criticize me, or even resort to ad hominem attacks.
Stratigraphy and sedimentology aren't relevant to the central question, even if I didn't know anything about them. And I do anyway. Still, you could try to educate us. What about stratigraphy, etc., renders the cryptic species problem nonexistent?
As far as I am concerned this discussion is now over.
...for the fifth time, by my count, though I may have missed one or two.
You certainly don't act like you do have a background in stratigraphy and sedimentology, even if you did take a course. Anyway, between you and I we have been replaying the debate on Punctuated Equilibrium which had Levinton and Schopf on one side, and Eldredge, Gould and Stanley on the other. I'm not interested in posting more "spilled ink" over this. Have a good day John. C'est fini.

John Harshman · 31 January 2011

John Kwok said: I'm not interested in posting more "spilled ink" over this.
No, we haven't been repeating a debate, since the original didn't involve, on one side, simple accusations of "you don't know stratigraphy".
Have a good day John. C'est fini.
6th time.

John Kwok · 31 January 2011

John Harshman said:
John Kwok said: I'm not interested in posting more "spilled ink" over this.
No, we haven't been repeating a debate, since the original didn't involve, on one side, simple accusations of "you don't know stratigraphy".
Have a good day John. C'est fini.
6th time.
You have not addressed any of my arguments with regards to stratigraphy and sedimentology, so I have to wonder whether you have a background. Anyway, if you read your history of science, you will see that similar - if not identical - arguments have been made for years. Where we might agree is realizing that Punctuated Equilbrium itself does not pose a direct challenge to the Modern Synthesis Theory of Evolution, though there are some who believe it does.

John Harshman · 31 January 2011

John Kwok said: You have not addressed any of my arguments with regards to stratigraphy and sedimentology, so I have to wonder whether you have a background.
You are correct that I have not addressed any of your stratigraphic arguments. In fact I have failed even to recognize that you were making any such arguments. Perhaps you could repeat them here, briefly, so I could try to at least notice them.
Anyway, if you read your history of science, you will see that similar - if not identical - arguments have been made for years.
Not sure which arguments you mean here, or what history of science you mean. Is there indeed a history of PE somewhere?
Where we might agree is realizing that Punctuated Equilbrium itself does not pose a direct challenge to the Modern Synthesis Theory of Evolution, though there are some who believe it does.
That depends on what you think the content of PE is. I think the original formulation does constitute a challenge to the Modern Synthesis, in that it requires a mechanism that prevents adaptive evolution except under special conditions (speciation in small, peripheral populations). Since we know of no such mechanism, acceptance of PE would suggest that the synthesis is be least very incomplete. Now, if you reduce PE to the idea that there is widespread stasis, that doesn't necessarily conflict at all.

John Kwok · 31 January 2011

John Harshman said:
John Kwok said: You have not addressed any of my arguments with regards to stratigraphy and sedimentology, so I have to wonder whether you have a background.
You are correct that I have not addressed any of your stratigraphic arguments. In fact I have failed even to recognize that you were making any such arguments. Perhaps you could repeat them here, briefly, so I could try to at least notice them.
Anyway, if you read your history of science, you will see that similar - if not identical - arguments have been made for years.
Not sure which arguments you mean here, or what history of science you mean. Is there indeed a history of PE somewhere?
Where we might agree is realizing that Punctuated Equilbrium itself does not pose a direct challenge to the Modern Synthesis Theory of Evolution, though there are some who believe it does.
That depends on what you think the content of PE is. I think the original formulation does constitute a challenge to the Modern Synthesis, in that it requires a mechanism that prevents adaptive evolution except under special conditions (speciation in small, peripheral populations). Since we know of no such mechanism, acceptance of PE would suggest that the synthesis is be least very incomplete. Now, if you reduce PE to the idea that there is widespread stasis, that doesn't necessarily conflict at all.
I have said more than once that if you have a stratigraphic section that shows continuous sedimentation and you sample broadly enough to rule out the arrival of immigrant taxa, then the lineage-splitting event is probably at the very least, a morphospecies speciation event. As for how you are reading Punctuated Equilibrium, even as far back as the early to mid 1970s, none other than Ernst Mayr regarded it as a simple extrapolation of allopatric speciation as applied to the fossil record (which, I might add, is what Eldredge himself was hinting at in his very first paper on it - approximately a year before the Eldredge and Gould 1972 paper which formally introduced Punctuated Equilibrium). Both points I have made more than once and you have either ignored them or dismissed them. Really have no more time to discuss this further.

John Harshman · 1 February 2011

John Kwok said: I have said more than once that if you have a stratigraphic section that shows continuous sedimentation and you sample broadly enough to rule out the arrival of immigrant taxa, then the lineage-splitting event is probably at the very least, a morphospecies speciation event.
Yes, you have said this more than once. And every time I have pointed out that this isn't true. You can't equate a morphospecies speciation event (which is nothing more than a difference between earlier and later populations) with a lineage-splitting event unless you make the assumption that no cryptic species are present. Continuous sedimentation is irrelevant. The very best record, even preservation of every individual in the populations, will not distinguish morphological change with lineage splitting from morphological change without splitting in one of a set of cryptic species.
As for how you are reading Punctuated Equilibrium, even as far back as the early to mid 1970s, none other than Ernst Mayr regarded it as a simple extrapolation of allopatric speciation as applied to the fossil record (which, I might add, is what Eldredge himself was hinting at in his very first paper on it - approximately a year before the Eldredge and Gould 1972 paper which formally introduced Punctuated Equilibrium).
Not just allopatric speciation: Mayr's particular model of allopatric speciation, or peripatric speciation. Mayr's model requires that most of the time, in large populations, the existence of "coadapted gene complexes" prevents response to selection. Only in small, isolated populations, in which drift can overcome selection, can a "genetic revolution" allow rapid change. This is the original mechanism of stasis and rapid change assumed by PE. We can now be pretty sure that Mayr's ideas were just wrong. PE is thus left with a choice: find a new mechanism by which stasis can be broken only during speciation (which is the challenge to the modern synthesis), or abandon claims to be a theory about speciation and just talk about stasis (my second option).
Both points I have made more than once and you have either ignored them or dismissed them. Really have no more time to discuss this further.
7th time. Yes, you make your points over and over, and over and over I explain why they're wrong, and you never respond except to repeat your original claim.

John Kwok · 1 February 2011

Sorry John, I strongly disagree with all of your points. You seem to think that cryptic species are far more common, when, having studied with David Jablonski, you should have garnered some insight with regards to the role of larval dispersal in affecting the survival of species, especially during mass extinctions (If we have widely distributed organisms that rely upon larval dispersal, then it wouldn't be possible to have sufficient reproductive isolation occurring that would result in cryptic species.). If cryptic species were really a problem, then I doubt that prominent neontologists like Douglas Futuyma and Massimo Pigliucci, would recognized the reality of evolutionary stasis.

The most interesting aspects of Punctuated Equilibrium are its recognition of evolutionary stasis and that species are real entities that have births and deaths. Not once have you addressed these.

Anyway this discussion has gone on far enough. I am still trying to fight off a bad cold and must return to my writing.

John Harshman · 1 February 2011

John Kwok said: Sorry John, I strongly disagree with all of your points.
I know you do. You just haven't, so far, bothered to explain why.
You seem to think that cryptic species are far more common, when, having studied with David Jablonski, you should have garnered some insight with regards to the role of larval dispersal in affecting the survival of species, especially during mass extinctions (If we have widely distributed organisms that rely upon larval dispersal, then it wouldn't be possible to have sufficient reproductive isolation occurring that would result in cryptic species.).
Yes, I'm familiar with Jablonski's work, but it's bizarre that you invoke it here. If anything, that's an argument against allopatric speciation -- any evolution of reproductive isolation -- not against cryptic species. So what you're saying is that groups with planktonic larva are unlikely to have a high speciation rate. But that says nothing about whether the few new species will be cryptic or otherwise.
If cryptic species were really a problem, then I doubt that prominent neontologists like Douglas Futuyma and Massimo Pigliucci, would recognized the reality of evolutionary stasis.
Again you misunderstand completely. First, this is just an argument from authority. Second, the argument isn't about stasis, but about the coincidence of punctuation with lineage splitting. So this "argument" is invalid on two fronts.
The most interesting aspects of Punctuated Equilibrium are its recognition of evolutionary stasis and that species are real entities that have births and deaths. Not once have you addressed these.
With good reason. I'm not arguing about those aspects. Stasis might in fact easily be a real and widespread phenomenon, and I have never been sure what it means for species to be "real entities". Now if it means there is a prevalence of sharp, near-instant "speciation events", as well as budding in which an ancestral species persists unchanged while spinning off a new one, I doubt that, though again that claim has no necessary connection to the central claim of PE, the coincidence of speciation and morphological change.
Anyway this discussion has gone on far enough. I am still trying to fight off a bad cold and must return to my writing.
8th time.

Malchus · 1 February 2011

The reason that Kwok has not actually addressed your points is, I suspect, because he can't. But John is quite capable of name-dropping in every post. Kwok, either explain why Harshman's responses are invalid or simply stop posting. You're making yourself look more and more as though you really don't have any good understanding of the point that Harshman is raising.
John Harshman said:
John Kwok said: I have said more than once that if you have a stratigraphic section that shows continuous sedimentation and you sample broadly enough to rule out the arrival of immigrant taxa, then the lineage-splitting event is probably at the very least, a morphospecies speciation event.
Yes, you have said this more than once. And every time I have pointed out that this isn't true. You can't equate a morphospecies speciation event (which is nothing more than a difference between earlier and later populations) with a lineage-splitting event unless you make the assumption that no cryptic species are present. Continuous sedimentation is irrelevant. The very best record, even preservation of every individual in the populations, will not distinguish morphological change with lineage splitting from morphological change without splitting in one of a set of cryptic species.
As for how you are reading Punctuated Equilibrium, even as far back as the early to mid 1970s, none other than Ernst Mayr regarded it as a simple extrapolation of allopatric speciation as applied to the fossil record (which, I might add, is what Eldredge himself was hinting at in his very first paper on it - approximately a year before the Eldredge and Gould 1972 paper which formally introduced Punctuated Equilibrium).
Not just allopatric speciation: Mayr's particular model of allopatric speciation, or peripatric speciation. Mayr's model requires that most of the time, in large populations, the existence of "coadapted gene complexes" prevents response to selection. Only in small, isolated populations, in which drift can overcome selection, can a "genetic revolution" allow rapid change. This is the original mechanism of stasis and rapid change assumed by PE. We can now be pretty sure that Mayr's ideas were just wrong. PE is thus left with a choice: find a new mechanism by which stasis can be broken only during speciation (which is the challenge to the modern synthesis), or abandon claims to be a theory about speciation and just talk about stasis (my second option).
Both points I have made more than once and you have either ignored them or dismissed them. Really have no more time to discuss this further.
7th time. Yes, you make your points over and over, and over and over I explain why they're wrong, and you never respond except to repeat your original claim.

Malchus · 1 February 2011

No, actually you haven't. It's becoming fairly clear that you simply don't understand the problem. You might try learning something about evolutionary biology, John, before jumping to an incorrect position on a point like this. I realize that biology isn't your field - computer programming, I believe is your area - but really. Evolutionary biology isn't that complicated.
John Kwok said:
Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.

John Kwok · 1 February 2011

1) No, if you have very patchy environments which foster reproductive isolation, you would still see speciation occurring. During times of normal, background extinction, those taxa that have broader larval dispersal, would be those least resistant to extinction (and the ones most likely to exhibit long-term evolutionary stasis). i would also suspect that given their broader geographic ranges, it would be hard to separate out cryptic species if they exist within.

2) Instead of arguing with me over the prevalence of evolutionary stasis, at least amongst marine metazoans, dig it up in the literature. That literature does exist. Again I'm not your online library resource.

John Kwok · 1 February 2011

Malchus said: No, actually you haven't. It's becoming fairly clear that you simply don't understand the problem. You might try learning something about evolutionary biology, John, before jumping to an incorrect position on a point like this. I realize that biology isn't your field - computer programming, I believe is your area - but really. Evolutionary biology isn't that complicated.
John Kwok said:
Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.
You're absolutely right, evolutionary biology isn't complicated at all. What is complicated however, are the perspectives of a neontologist (Harshman) who has worked only with vertebrates and a former invertebrate paleobiologist (yours truly).

John Kwok · 1 February 2011

John Kwok said: 1) No, if you have very patchy environments which foster reproductive isolation, you would still see speciation occurring. During times of normal, background extinction, those taxa that have broader larval dispersal, would be those least resistant to extinction (and the ones most likely to exhibit long-term evolutionary stasis). i would also suspect that given their broader geographic ranges, it would be hard to separate out cryptic species if they exist within. 2) Instead of arguing with me over the prevalence of evolutionary stasis, at least amongst marine metazoans, dig it up in the literature. That literature does exist. Again I'm not your online library resource.
And John, I'm not disputing that there are cryptic species have been present in the history of life prior to the Recent. But all we have for the last five hundred forty-three odd million years is a fossil record based on hard-part shelly invertebrates, and I have to use that data, nothing else, to support the prevalence of morphological stasis at least amongst Marine metazoa. If you want a relatively recent example of some kind of long-term stasis, may I suggest looking at the work published by Derek Briggs and his team (I don't think he's the lead author BTW) that was published last spring in Nature on the discovery of a Burgess Shale-like fauna in Early Ordovician strata from North Africa. It's evidence like that I base my conclusions, not what Eldredge, Gould, Futuyma or Pigliucci have said.

John Kwok · 1 February 2011

And you Malchus seem incapable of doing anything but hurtling insults at me. Have yet to read one substantial contribution you have tried to make with regards to either sedimentology geology or evolutionary biology in reply to my comments in this thread.

The existence of morphological stasis in the fossil record is something that invertebrate paleontologists in the 19th Century recognized. However, it wasn't until the advent of the Modern Synthesis that we had a younger generation of inverebrate paleobiologists like Eldredge, Gould, Stanley and others, who opted to gain insights from the Modern Synthesis into explaining patterns such as morphological stasis that they were seeing in the fossil record (Though in this case, paleobiology came late to the "game" so to speak, and it was only due to the "Young Turks" like those I have cited.).

As for myself, I was never really interested in determining which mode of speciation was responsible for punctuated equilibrium. Instead, what I found of interest was the long-term persistence of morphologically identical taxa over time, especially the long-term survival of taxa existing in the same, or similar, enviroments, which have been the subject of extensive research by some University of Chicago paleobiologists and their former students.

John Harshman · 1 February 2011

John Kwok said: 1) No, if you have very patchy environments which foster reproductive isolation, you would still see speciation occurring. During times of normal, background extinction, those taxa that have broader larval dispersal, would be those least resistant to extinction (and the ones most likely to exhibit long-term evolutionary stasis). i would also suspect that given their broader geographic ranges, it would be hard to separate out cryptic species if they exist within.
No to what? I fail to see the relevance of brooding vs. planktonic larvae to anything we've been talking about.
2) Instead of arguing with me over the prevalence of evolutionary stasis, at least amongst marine metazoans, dig it up in the literature. That literature does exist. Again I'm not your online library resource.
I'm not arguing with you over the prevalence of evolutionary stasis. I'm arguing with you about our ability to recognize splitting events in the fossil record. (I also question whether the case for stasis as a dominant phenomenon has been adequately made, but that isn't what we've been discussing here.) It's amazing to me that after all this time you still don't know what we're arguing about.
And John, I'm not disputing that there are cryptic species have been present in the history of life prior to the Recent. But all we have for the last five hundred forty-three odd million years is a fossil record based on hard-part shelly invertebrates, and I have to use that data, nothing else, to support the prevalence of morphological stasis at least amongst Marine metazoa.
We aren't arguing about morphological stasis. Are you on drugs? (Not out of the question, given your cold.)
If you want a relatively recent example of some kind of long-term stasis, may I suggest looking at the work published by Derek Briggs and his team (I don't think he's the lead author BTW) that was published last spring in Nature on the discovery of a Burgess Shale-like fauna in Early Ordovician strata from North Africa. It's evidence like that I base my conclusions, not what Eldredge, Gould, Futuyma or Pigliucci have said.
I recall the paper. But are the morphospecies in that fauna the same morphospecies as those in the Burgess? That, I don't recall. What percentage of described species are common between faunas? Genera don't count; stasis is a phenomenon within morphospecies. But again, we aren't arguing about stasis and never have been. Remember that.

John Kwok · 1 February 2011

1) I do in the sense that those taxa that are geographically more broadly distributed - as a result of their laval dispersal strategies - would be the ones most likely to exhibit prolonged morphological stasis.

2) I think we can recognize splitting events in marine Metazoans. Where we can't recognize them obviously is in terrestrial environments since these aren't nearly as well-preserved as nearshore marine ones.

3) No, I believe they are not the same species, but instead, belong to the same clades as the Burgess Shale ones. If you want relevant examples, I suggest you look at Brett et al.'s work for starters. And now you seem to accept morphological stasis as a reality, when for the longest time you were questioning its existence.

John Kwok · 1 February 2011

John Kwok said: 1) I do in the sense that those taxa that are geographically more broadly distributed - as a result of their laval dispersal strategies - would be the ones most likely to exhibit prolonged morphological stasis.
And even what I wrote (above) may not be true with taxa possessing plantonic larvae if there are reproductive isolating mechanisms such as long-term changes in the direction and intensity of currents like the Gulf Stream, for example, changes in water temperature that may or may not be due to changes in currents, or the erection of unexpected barriers due to, for example, an extremely active period of volcanism that could create barriers such as islands or isthmuses which would result in dividing one widespread population into two.

John Harshman · 1 February 2011

John Kwok said: 1) I do in the sense that those taxa that are geographically more broadly distributed - as a result of their laval dispersal strategies - would be the ones most likely to exhibit prolonged morphological stasis.
You do what? And I repeat that we are not arguing about stasis, and also my question of whether you are on drugs.
2) I think we can recognize splitting events in marine Metazoans. Where we can't recognize them obviously is in terrestrial environments since these aren't nearly as well-preserved as nearshore marine ones.
What does quality of preservation have to do with it, and why do you think we can recognize splitting events in marine metazoans?
3) No, I believe they are not the same species, but instead, belong to the same clades as the Burgess Shale ones. If you want relevant examples, I suggest you look at Brett et al.'s work for starters. And now you seem to accept morphological stasis as a reality, when for the longest time you were questioning its existence.
If they don't belong to the same species, that isn't stasis, at least by the definition of PE. Clades can encompass any desired amount of differentiation. Of course they belong to the same clades. All life belongs to the same clades. It isn't that I doubt the reality of stasis. What I doubt is its ubiquity. And I repeat that that has never been what we were arguing about. I continue to marvel at your inability to understand this simple fact.

Shebardigan · 1 February 2011

John Harshman said: Or, if your cold and your novel are too pressing, just stop responding.
Kount Kwokula has remarkable recuperative powers. Would they were accompanied with other capabilities not, unfortunately, in evidence.

Malchus · 1 February 2011

Kwok, my point is mote basic: I am not responding to your sedimentary points because they are not relevant to the point that Harshman is making. He is right - you really don't have any idea what the discussion is about. No one is disputing morphological stasis. No one. But it's in any way relevant to Harshman's point. Kwok, if you're that confused and distracted, you should simply admit your error and stop arguing from authority on a point you don't even understand.
John Kwok said: And you Malchus seem incapable of doing anything but hurtling insults at me. Have yet to read one substantial contribution you have tried to make with regards to either sedimentology geology or evolutionary biology in reply to my comments in this thread. The existence of morphological stasis in the fossil record is something that invertebrate paleontologists in the 19th Century recognized. However, it wasn't until the advent of the Modern Synthesis that we had a younger generation of inverebrate paleobiologists like Eldredge, Gould, Stanley and others, who opted to gain insights from the Modern Synthesis into explaining patterns such as morphological stasis that they were seeing in the fossil record (Though in this case, paleobiology came late to the "game" so to speak, and it was only due to the "Young Turks" like those I have cited.). As for myself, I was never really interested in determining which mode of speciation was responsible for punctuated equilibrium. Instead, what I found of interest was the long-term persistence of morphologically identical taxa over time, especially the long-term survival of taxa existing in the same, or similar, enviroments, which have been the subject of extensive research by some University of Chicago paleobiologists and their former students.

Malchus · 1 February 2011

Kwok, you were never an invertebrate paleobiologist. As I recall, you took a few courses in the field. Exaggerating your accomplishments wins you no points here.
John Kwok said:
Malchus said: No, actually you haven't. It's becoming fairly clear that you simply don't understand the problem. You might try learning something about evolutionary biology, John, before jumping to an incorrect position on a point like this. I realize that biology isn't your field - computer programming, I believe is your area - but really. Evolutionary biology isn't that complicated.
John Kwok said:
Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.
You're absolutely right, evolutionary biology isn't complicated at all. What is complicated however, are the perspectives of a neontologist (Harshman) who has worked only with vertebrates and a former invertebrate paleobiologist (yours truly).

John Kwok · 2 February 2011

Malchus said: Kwok, you were never an invertebrate paleobiologist. As I recall, you took a few courses in the field. Exaggerating your accomplishments wins you no points here.
John Kwok said:
Malchus said: No, actually you haven't. It's becoming fairly clear that you simply don't understand the problem. You might try learning something about evolutionary biology, John, before jumping to an incorrect position on a point like this. I realize that biology isn't your field - computer programming, I believe is your area - but really. Evolutionary biology isn't that complicated.
John Kwok said:
Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.
You're absolutely right, evolutionary biology isn't complicated at all. What is complicated however, are the perspectives of a neontologist (Harshman) who has worked only with vertebrates and a former invertebrate paleobiologist (yours truly).
I have a master's degree in which my specialty was invertebrate paleobiology and I reviewed several manuscripts that were published in several notable scientific journals. That's more than I can say about you. You'll have to excuse me, but I'm trying to shake off a very, very bad cold.

ben · 2 February 2011

I propose Ben's Law:

"Given enough time, every conversation which includes John Kwok will eventually become about John Kwok."

mrg · 2 February 2011

ben said:[SNIP}
"Eventually"?

ben · 2 February 2011

mrg said:
ben said:[SNIP}
"Eventually"?
I also propose Trombley's Corollary* to Ben's Law: "Any conversation which includes John Kwok, regardless of duration, is already thought by John Kwok to be about John Kwok." ----- *Named after Mike Trombley, former major league baseball pitcher and 1985 Western Massachusetts high school football Superbowl-winning quarterback who happens to have been a classmate of mine at Minnechaug Regional High School in Wilbraham Massachusetts (also home, incidentally, to the Friendly's ice cream corporation) and who I sat next to in Advanced Writing class in 12th grade, often helping the good-natured but not overly bright Mr Trombley with classwork, which direct and meaningful relationship is the reason I named the corollary after him instead of former Notre Dame and San Francisco 49ers quarterback, three-time Super Bowl MVP and NFL hall of famer Joe Montana, to whom (with his lovely wife and children at his side) I once served a peach-flavored Snapple iced tea, for which he generously allowed me to keep the seventy cents change on the $1.30 purchase, or after Far Rockaway, Queens (which I have never visited, though I have been a few times to Manhattan, home of, gosh, so many famous people that I could namedrop for hours were I not motivated to condense this current thought into the most compact and efficient communication possible) native Richard Feynman, the Nobel Prize-winning physicist and member of the Rogers Commission (which investigated the explosion of the space shuttle Challenger which, incidentally, occurred on my first day of college at the prestigious University of Massachusetts where my wife happens to have once attended an Education class with current NBA star and 2007 NBA defensive player of the year Marcus Camby) who seems to have been a really interesting person who I never met or had anything to do with but I am dropping his name here just because and if you have a problem with that you owe me a new set of snow tires and a pair of those little hydraulic cylinders that hold the back hatch open on my 1999 Saab 9-3 (base model) with 145,000 (mostly highway) miles, a persistent check engine light and an annoying body squeak when you go over bumps in cold weather, really a nice car but a little over-engineered and expensive to fix, it's probably just a matter of time before something breaks which in a Ford or Toyota would be an affordable repair but in the Saab would be too expensive to be worth it.

Wolfhound · 2 February 2011

LOL!

Malchus · 6 February 2011

In other words, you admit that you were never an invertebrate paleobiologist. You were merely a student - it wasn't even the subject of your masters. And you've no idea what my field is. Really, Kwok, grow up.
John Kwok said:
Malchus said: Kwok, you were never an invertebrate paleobiologist. As I recall, you took a few courses in the field. Exaggerating your accomplishments wins you no points here.
John Kwok said:
Malchus said: No, actually you haven't. It's becoming fairly clear that you simply don't understand the problem. You might try learning something about evolutionary biology, John, before jumping to an incorrect position on a point like this. I realize that biology isn't your field - computer programming, I believe is your area - but really. Evolutionary biology isn't that complicated.
John Kwok said:
Malchus said: And I might add that your counter-point does not actually address Harshman's argument. Apparently you are unable to do so.
It does Malchus but you are simply too dense to realize it. I suggest you delve into the literature and learn something about stratigraphy and sedimentology. As for not addressing Harshman's points, that's baloney. I have, but he doesn't like my answers.
You're absolutely right, evolutionary biology isn't complicated at all. What is complicated however, are the perspectives of a neontologist (Harshman) who has worked only with vertebrates and a former invertebrate paleobiologist (yours truly).
I have a master's degree in which my specialty was invertebrate paleobiology and I reviewed several manuscripts that were published in several notable scientific journals. That's more than I can say about you. You'll have to excuse me, but I'm trying to shake off a very, very bad cold.

Dale Husband · 6 February 2011

Malchus/John Kwok said: [A lot of pointless griping about trivial differences.]
SHUT THE HELL UP!