"Intraspecific macroevolution" within domestic dog breeds

Posted 29 July 2010 by Nick Matzke

Several years ago, I saw a fantastic talk at the Evolution meeting about Intraspecific macroevolution: variation of cranial shape in dog breeds. The talk was by Abby Drake, then a grad student, and reported on a huge digital morphometric comparison of the skulls of dogs and many representatives from the order Carnivora (dogs, cats, bears, sea lions, etc.). Morphometrics basically consists of taking digital photos of e.g. bones from different angles, and then marking the same landmarks on homologous bones across a big group. Then you can quantitatively compare the differences in shape, independent of things like body size. This is a much more sophisticated analysis than is possible with just calipers, where you can only get length, width, etc. A previous study had noted that the skull variation in dogs was bigger than the variation in the family Canidae, but the incredible result of Drake's study was that the variation in shape of dog skulls was bigger than the variation in shape across the entire order Carnivora, which is 60 million years old and includes even mostly-aquatic forms.

Figure 3: Principal component (PC) analysis for skull shape in the complete data set. A-C, Plots of the PC scores. D, Shape changes associated with the PC axes. For each PC, the shapes corresponding to the observed extremes in the positive and negative directions are shown as a warped surface of a wolf skull (Wiley et al. 2005).

And most of this morphological variation took only a few hundred years to produce. It is true that some of these weird skulls would not be favored in the wild -- Drake notes that natural selection is reduced when your food comes from a can rather than stuff you hunt -- and artificial selection is greatly enhanced by selective breeding. But this is strong evidence that (a) there is no problem on the genetic variability end of the equation for the kinds of variability that we see in a mammalian order like Carnivores; rather the constraint is natural selection for a particular niche. If the selective pressure is there, the morphological change can happen very quickly; and (b) lack of time isn't the issue; if the conditions are right, hundreds or thousands of years can be plenty of time. I didn't even notice when this study came out and got a bunch of press in January, probably because I actually had a girlfriend at the time (see, rare events do happen in geologic time!). But this is a study that should be in the back pocket of any creationism opponent. You can see an example of its usage on Cornelius Hunter here; it's kind of like a surprise sack of a quarterback. The other thing I like about the conclusion of "intraspecific macroevolution" is that it tweaks a lot of standard tropes that even we scientists have about what is meant by the word "macroevolution." The minimal definition of macroevolution is "evolution above the species level", but it has become a catchall term encompassing everything from speciation to lineage-diversification and extinction dynamics to "evolution of 'higher taxa'" (ack! go read "down with phyla!") to vaguely defined "large" amounts of change to evo-devo changes in development. These things then all get mixed together in people's heads, resulting in the erroneous presumption that "'large' amounts of change" = lots of speciation events = the origin of some big Linnaean 'taxon' = lots of action at the lineage-counting level. As a very rough approximation it might be true that these different things are often linked, but as this study shows, it ain't always true. We would probably be better off using specific terms for each of these different topics, and not trying to lump them all together under "macroevolution" as if they were all intrinsically connected. Questions like "is macroevolution just the result of repeated rounds of microevolution" have almost no meaning if "macroevolution" refers to all of these different things at once. References

Abby Grace Drake and Christian Peter Klingenberg Large‐Scale Diversification of Skull Shape in Domestic Dogs: Disparity and Modularity Am Nat 2010. Vol. 175, pp. 289-301 DOI: 10.1086/650372 Abstract: The variation among domestic dog breeds offers a unique opportunity to study large‐scale diversification by microevolutionary mechanisms. We use geometric morphometrics to quantify the diversity of skull shape in 106 breeds of domestic dog, in three wild canid species, and across the order Carnivora. The amount of shape variation among domestic dogs far exceeds that in wild species, and it is comparable to the disparity throughout the Carnivora. The greatest shape distances between dog breeds clearly surpass the maximum divergence between species in the Carnivora. Moreover, domestic dogs occupy a range of novel shapes outside the domain of wild carnivorans. The disparity among companion dogs substantially exceeds that of other classes of breeds, suggesting that relaxed functional demands facilitated diversification. Much of the diversity of dog skull shapes stems from variation between short and elongate skulls and from modularity of the face versus that of the neurocranium. These patterns of integration and modularity apply to variation among individuals and breeds, but they also apply to fluctuating asymmetry, indicating they have a shared developmental basis. These patterns of variation are also found for the wolf and across the Carnivora, suggesting that they existed before the domestication of dogs and are not a result of selective breeding.

Robert K. Wayne (1986). "Cranial Morphology of Domestic and Wild Canids: The Influence of Development on Morphological Change." Evolution, Vol. 40, No. 2 (Mar., 1986), pp. 243-261 * Stable URL: http://www.jstor.org/stable/2408805 Abstract l The domestic dog varies remarkably in cranial morphology. In fact, the differences in size and proportion between some dog breeds are as great as those between many genera of wild canids. In this study, I compare patterns of intracranial allometry and morphologic diversity between the domestic dog and wild canid species. The results demonstrate that the domestic dog is morphologically distinct from all other canids except its close relatives, the wolf-like canids. Following this, I compare patterns of static and ontogenetic scaling. Data on growth of domestic dogs are presented and used to investigate the developmental mechanisms underlying breed evolution. Apparently, most small breeds are paedomorphic with respect to certain morphologic characters. In dogs and other domestic animals, morphologic diversity among adults seems to depend on that expressed during development.

66 Comments

Steve · 29 July 2010

"if the conditions are right, hundreds or thousands of years can be plenty of time."

Yup, and I predict it will be a matter of days before the creationists twist this study to claim it proves that the few "kinds" on the Ark could easily be pushed (by God, natch) to fill all the niches with all the different species we see today.

But then, for creationists, the conditions are always right - for lying.

TR Gregory · 29 July 2010

“intraspecific macroevolution”

Cringe. This is an oxymoron. Plus, the morphological differences within this species, large as they are, are not permanent at all. They are easily lost when artificial genetic isolation is relaxed for even a couple of generations. Need data, sloppy terminology.

TR Gregory · 29 July 2010

*neat data

Nick (Matzke) · 29 July 2010

Hi Ryan! Thanks for the comment, I was attempting to be provocative (obviously). So I'll continue:

1. Who says these aren't permanent changes? And, who says that the morphological changes that *do* occur in "regular large scale natural evolution" *are* permanent? In either case what happens morphologically probably depends substantially on the future selective regime, which will either maintain the changes, or it won't.

2. Re: oxymoron. So, hypothetically, if a major developmental change leading to a "major" morphological change (say, on the level of the differences we typically see between genera or between families) did occur *within* a species, complete with polymorophism within the population for this change which is later fixed by selection -- but with no speciation/lineage-splitting event, would this be:

(a) a microevolutionary event, because it was within a species, or

(b) a macroevolutionary event, because the amount of change was on the level of the differences often observed between higher taxa?

Obviously there must be a simple, correct answer, right?

;-)

JGB · 29 July 2010

TR Gregory said: “intraspecific macroevolution” They are easily lost when artificial genetic isolation is relaxed for even a couple of generations.

In that sense and in light of our partial Neanderthal ancestry what major changes (morphological or not) might just as easily have been lost when if that genetic isolation was lost for longer? In other words what is the significance of emphasizing the incomplete speciation of the different dog breeds?

MrG · 29 July 2010

Steve said: Yup, and I predict it will be a matter of days before the creationists twist this study ...

Though I may be overly suspicious here, I think your timescale may have been optimistic:

TR Gregory said: “intraspecific macroevolution” Cringe. This is an oxymoron.

Fascinating article, NickM. The variations in morphology among dogs are obvious, but that they exceed by proper measures those of the rest of the Carnivora comes as a surprise.

DS · 29 July 2010

BUT THEY ARE ALL STILL DOGS!

Well, somebody had to say it.

The important thing is not the degree of morphological or even genetic variation. The important thing is genetic discontinuity. Now if all of the types of dogs are still potentially interbreeding, then there may or may not be any significant discontinuities. That would depend on the actual degree of interbreeding, along with mutation rate, population size, epistatic interactions, etc. Given the degree of variation observed, speciation would seem to be a fairly trivial outcome of such processes. This does demonstrate the tremendous power of artificial selection. It should also convince any unbiased observer that natural selection can be a very powerful force as well.

Dave C · 29 July 2010

MrG said:
Steve said: Yup, and I predict it will be a matter of days before the creationists twist this study ...
Though I may be overly suspicious here, I think your timescale may have been optimistic:
TR Gregory said: “intraspecific macroevolution” Cringe. This is an oxymoron.
Fascinating article, NickM. The variations in morphology among dogs are obvious, but that they exceed by proper measures those of the rest of the Carnivora comes as a surprise.

You are indeed being overly suspicious. Dr. Gregory is an evolutionary biologist who literally wrote the book on genome evolution (full disclosure: I haven't read it). Definitely not a creationist.

MrPeach · 29 July 2010

As I read this article about humans making huge changes in the morphological diversity of a companion specie, I could not help but speculate as to how we have guided our OWN morphological changes. Anyone have any thoughts?

Dave Luckett · 30 July 2010

I would not care to put money on the long survival of male facial and body hair, for one thing.

Mike Clinch · 30 July 2010

TR Gregory said: “intraspecific macroevolution” Cringe. This is an oxymoron. Plus, the morphological differences within this species, large as they are, are not permanent at all. They are easily lost when artificial genetic isolation is relaxed for even a couple of generations. Need data, sloppy terminology.

I'm not a biologist (glacial geologist) but "intraspecific macroevolution" looks like a loser to me. Any creationist or ID'er (except a diehard YEC) will immediately dismiss the paper as demonstrating "microevolution" that has no bearing on "macroevolution", while the YEC will blather on about how rapidly the "kinds" will change to repopulate the earth after Noah's Flood. To me, it's just evolution. The lesson of this paper is that the dog genome is capable of producing more than sufficient variability in body size and morphology in a very short period of time. The variability is severely limited by natural selection in the wild, leading to a much more restricted morphology in wild dogs, and reversion to this type in feral dog communities. The variability is preserved by dog breeders, or even facilitated by them. In the long run, we might even suggest that the dog species has already "evolved" into several subspecies or new species. After all, if a species is defined as a population that interbreds and produces viable offspring, I find it hard to believe that a Chihuahua and a St. Bernard can be defined as the same species. To me, the utility of this paper is to point out the speed of change in the genome of a single species in situations where the variability can be preserved. Such might be the case after mass extinctions, where lack of competition and the emptying of most ecological niches have led to explosive radiation of mammals in the early Tertiary, of dinosaurs in the early Mesozoic, and of many other organisms. Creationists claim that there is no mechanism for the recoveries after mass extinctions. This paper provides one - normal evolution.

John Kwok · 30 July 2010

Sorry Nick, but morphometrics is similar the mathematics of body size and shape. While today virtually all morphometric data capture is via digitizing of images, it started with the use of calipers. After one collects the data, and then (hopefully) plots it just to see the amount of dispersion in it, then what is done next is usually some kind of data reduction technique to see if there are any correlations between body size and shape or unseen factors which account for it. What is normally referred to as factor analysis - if it is unrotated - is merely principal component analysis and this is the technique most widely used for data reduction of morphometric data (Stephen Jay Gould has an elegant description of principal components analysis - which if my memory serves, he refers to incorrectly as "factor analysi" - in his "The Mismeasure of Man".).

John Kwok · 30 July 2010

Let me just rephrase the first sentence: Sorry Nick, but morphometrics is simply the mathematics of body size and shape.

John Kwok said: Sorry Nick, but morphometrics is similar the mathematics of body size and shape. While today virtually all morphometric data capture is via digitizing of images, it started with the use of calipers. After one collects the data, and then (hopefully) plots it just to see the amount of dispersion in it, then what is done next is usually some kind of data reduction technique to see if there are any correlations between body size and shape or unseen factors which account for it. What is normally referred to as factor analysis - if it is unrotated - is merely principal component analysis and this is the technique most widely used for data reduction of morphometric data (Stephen Jay Gould has an elegant description of principal components analysis - which if my memory serves, he refers to incorrectly as "factor analysi" - in his "The Mismeasure of Man".).

TomS · 30 July 2010

If I were a creationist, I would point out that this is confirmation that "intelligent design", in the form of human intervention, is capable of producing "macroevolution".

MrG · 30 July 2010

TomS said: If I were a creationist, I would point out that this is confirmation that "intelligent design", in the form of human intervention, is capable of producing "macroevolution".

Wot of it? Creationists are NEVER at a loss for a snappy answer. The nice thing about baloney is that the cost is extremely low and the supply indefinitely great.

eric · 30 July 2010

TomS said: If I were a creationist, I would point out that this is confirmation that "intelligent design", in the form of human intervention, is capable of producing "macroevolution".

Except that dog breeding occurred long before we could directly manipulate genomes. Dog breeding is therefore not intelligent design, its intelligent selection. The fact is, all the variation you see was produced by nature via either sexual reproductions standard mixing or mutation. We merely decided which variant bred with which. The fact that intelligent breeding can produce such rapid changes in morphology is, as Nick says, proof that its selection, not availability of variation, that keeps evolution relatively slow. Of course, if you were a creationist you'd probably ignore any counter-argument posted...

Wheels · 30 July 2010

Dogs appear to be one of the most morphologically plastic animals we've been able to domesticate. It seems like there's something unique about their genes to let us change their size, fur, body type, skull shape, and even innate behaviors to such a great degree. (Of course it could just be that we haven't domesticated rats for anything like the same length of time). I've heard that there's a single gene that's mostly responsible for the different angle of dog snouts (pointing out, up, or down compared to the rest of the skull), but haven't looked into that.

Tangentially related, but to me dogs always presented a ready challenge to the conventional definition of "species" anyway, since they interbreed so readily with wolves and coyotes (who also interbreed with each other) and produce fertile offspring.

MrG · 30 July 2010

Wheels said: Of course it could just be that we haven't domesticated rats for anything like the same length of time.

Points taken, but I would think that the case. Domestic pigeons get pretty wild, ditto for goldfish -- double tails, bubble eyes, even (I recollect) translucent goldfish where you can see their heart beating. Translucent ... try THAT with a dog.

MrG · 30 July 2010

eric said: Except that dog breeding occurred long before we could directly manipulate genomes. Dog breeding is therefore not intelligent design, its intelligent selection. The fact is, all the variation you see was produced by nature via either sexual reproductions standard mixing or mutation. We merely decided which variant bred with which.

Some of those selections are pretty weird (pekinese for example). As has been noted, I vaguely recall by Darwin himself in terms of pigeons, many of the "sports" of domesticated organisms are just something strange that popped up and breeders ran with. Also ... it is very difficult to say that the domestication of the wolf wasn't at the wolf's initiative. Those with a tame inclination were much more inclined to being domesticated than those that weren't.

Mike Elzinga · 30 July 2010

Wheels said: Dogs appear to be one of the most morphologically plastic animals we've been able to domesticate.

I suspect that this is what cats think about humans. ;-)

MrG · 30 July 2010

Mike Elzinga said: I suspect that this is what cats think about humans. ;-)

Actually, SCIENTIFIC AMERICAN did an article on cat descent and domestication that pretty much said cats domesticated US. I ran an outline in my blog, I'll give links if anyone cares. GIRL GENIUS fans of course think of Krosp, the Emperor of Cats: "I accept your fealty, human!"

Wheels · 30 July 2010

MrG said: Points taken, but I would think that the case. Domestic pigeons get pretty wild, ditto for goldfish -- double tails, bubble eyes, even (I recollect) translucent goldfish where you can see their heart beating.

True, good examples. As a counterpoint, domestic cats don't have nearly as much variety of basic shape built in it seems. Sure you get flat-faced Persians and adorable Scottish Folds, but beyond that cats still basically look like cats. You don't see cats bred to have limbs like their dorsal fins totally absent, or stumpy badger-hunting cats with high-set, short legs and down-pointing schnauses. I'm not sure this can be fully explained by lack of trying on the part of the sport breeding community.
Humans are so weird, taking perfectly functional wildlife and churning out mutant freaks like some kind of mad science, sometimes for no reason other than to do it. Not besmirching the habit*, just making an observation.
*Although I would like to see an end to breeding programs that cause obvious health problems, like Persians that can't stop "crying" or bulldogs that can barely breath.

MrG · 30 July 2010

Wheels said: True, good examples. As a counterpoint, domestic cats don't have nearly as much variety of basic shape built in it seems.

Yeah, the article discussed that, saying that the genetic variation between cats is roughly like IIRC French and Italians. But this was attributed partly to the fact that, other than mousing, cats aren't put to much work and so there's not so much reason to come up with specialized variants. And that's changing now due to artificial insemination, producing hybrids like the "caracat" (housecat-caracal hybrid).

Wheels · 30 July 2010

So the morphological variation in dogs mainly comes down to the fact that they can be put to some new use, uses more apparent and achievable to early breeders that weren't in this for sports, more than to some kind of easy plasticity built into their genome.

Mike Elzinga · 30 July 2010

MrG said: Yeah, the article discussed that, saying that the genetic variation between cats is roughly like IIRC French and Italians. But this was attributed partly to the fact that, other than mousing, cats aren't put to much work and so there's not so much reason to come up with specialized variants.

Dogs have masters; cats have staff.

MrG · 30 July 2010

Dogs come when called. Cats tell you to leave a message and they'll get back to you later.

Jedidiah Palosaari · 30 July 2010

It seems that this study would also indicate that when we identify different species by fossil evidence alone, we may artificially magnify the differences, and great diversity in one species may be mistaken as many different species. Would this be accurate?

John Kwok · 30 July 2010

I'm actually dealing with this issue for someone now, ghost-writing and editing a manuscript, but can't say anything further on that. But you have the right idea:

Jedidiah Palosaari said: It seems that this study would also indicate that when we identify different species by fossil evidence alone, we may artificially magnify the differences, and great diversity in one species may be mistaken as many different species. Would this be accurate?

eric · 30 July 2010

Jedidiah Palosaari said: It seems that this study would also indicate that when we identify different species by fossil evidence alone, we may artificially magnify the differences, and great diversity in one species may be mistaken as many different species. Would this be accurate?

Speciation being just the end result of variation, it would be fair to say that it might be dificult to tell the differences between variations in a widely varying species and species that have recently split. With some caveats. First lets get teh stupid out of the way. You understand that a continuum in morphology is a prediction of common descent with modification, and observing it is a confirmation of the theory of evolution, right? There is no other hypothesis or theory that rationally makes this prediction. 'Goddidit that way because he wanted to' certainly doesn't count...are we agreed? Okay, second: "by fossil evidence alone" is rarely the case, and even when it is, biologists consider more than just skull morphology. There are many other ways, for instance, to tell a wolf from a dog which make it quite clear which species is which. Lastly, biologists argue over what counts as a species in the first place. If you consider populations that don't breed with each other due to behavioral or geographical barriers (rather than biological barriers) to be different species, then of course some different species are going to have extremely similar physical traits (at least at first). So part of the 'mistakenness' depends on how you define species. The difficulty of distinguishing 'close species' versus 'wide variations' is, again, a problem one would predict given descent with modification.

Henry J · 30 July 2010

To put that in mathematical terms, sameness of species is not a transitive relation (i.e., A is like B and B is like C does not necessarily imply A is like C).

Nick (Matzke) · 30 July 2010

Jedidiah Palosaari said: It seems that this study would also indicate that when we identify different species by fossil evidence alone, we may artificially magnify the differences, and great diversity in one species may be mistaken as many different species. Would this be accurate?

Tough to say. Morphological change is only a very rough "clock" of "time since divergence". Sometimes you can have large change with low divergence time (e.g. dogs), other times you can have long divergence time with little morphological change (lots of examples, e.g. cryptic species). Based on the fact that with modern wild "species", critters that are skeletally almost indistinguishable often get split into different "species" on the basis of plumage, genetic divergence, etc., I think most paleontologists would say that "morphospecies" probably are a minimum estimate of the number of species existing at any given time. There are, though, cases where 2 different named fossil species turned out to be males and females of one species, or different developmental stages, etc.

Nick (Matzke) · 30 July 2010

John Kwok -- thanks, you're right I imagine, I haven't studied the history of this. However I doubt that anyone ever calipered all of the landmark-to-landmark distances, with 40 landmarks this would be 40^2 measurements per specimen. Easy with the computer, impossible without. Cheers!
Nick

Paul Burnett · 30 July 2010

Wheels said: Dogs appear to be one of the most morphologically plastic animals we've been able to domesticate.

Possibly, but go to a pet shop and look at goldfish. Or even look at the pigeons that Darwin wrote about.

John Kwok · 30 July 2010

Nick, With the latest digitizing equipment, it's really a relative piece of cake to collect morphometric data. But people were able to use calipers and, if I'm not mistaken, Stephen Jay Gould had published extensively in the mid to late 1960s and early 1970s using a lot of morphometric data derived from calipers. Ask Kevin Padian since I am sure he might steer to some useful histories on morphometric analysis. Best, John

Nick (Matzke) said: John Kwok -- thanks, you're right I imagine, I haven't studied the history of this. However I doubt that anyone ever calipered all of the landmark-to-landmark distances, with 40 landmarks this would be 40^2 measurements per specimen. Easy with the computer, impossible without. Cheers! Nick

Wheels · 30 July 2010

Dammit Nick, now I'm arguing with Cornelius Hunter again.
Why oh why couldn't it have been Yukon Cornelius instead?

Hygaboo Andersen · 31 July 2010

Gee, it's hard to think of a more rancid combination of arrogance and greed than the evolutionists when it comes to their pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature. Not only do they charge megabucks for 24-hour access to their articles, they also demand you register with them for the privilege of sending them this money! Evolutionists are worse than the pornographers their faith spawns when it comes to their money-grubbing spam machine! Indeed, what is the point in citing this crud knowing Christians will be unable to read and critique it--or is that the whole point?!

By the way, computer models of animals morphing into each other is not proof that it happened in real life! That is all this "research" really seems to amount to.

JGB · 31 July 2010

"pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature"

Apparently you ignored the ten years or so of acid test peer-review and debate the theory went through when we had the least amount of data when the vast majority of established scientists were opposed to the idea.

Or the 50 years of peer-review and commentary after that against a wide variety of other evolutionary theories radically different from Darwin's basic framework.

"citing this crud knowing Christians will be unable to read and critique it–or is that the whole point"

Well a simple google scholar search can uncover vast swaths of freely available literature for download. Most public universities libraries are very easy to access and free to read and review materials in the library without need of being a student. I could keep going with other ways that anybody can access this information, but I think you get the point

MrG · 31 July 2010

HA's just yanking your chain, JGB. "Anything for a laff."

Stanton · 31 July 2010

I would recommend some of my friends and contacts who help me get copies of pay-per-view reports, but, Hygaboo comes off as the sort of lazy fundamentalist ass-troll who regards reading as a hellbound blasphemy deserving of death, even if it's reading "See Spot Run" to children.

Stanton · 31 July 2010

JGB said:
pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature
Apparently you ignored the ten years or so of acid test peer-review and debate the theory went through when we had the least amount of data when the vast majority of established scientists were opposed to the idea. Or the 50 years of peer-review and commentary after that against a wide variety of other evolutionary theories radically different from Darwin's basic framework.

What did you expect for a fundamentalist troll? Reasoned, sincere thoughts or rudimentary social skills?

DS · 31 July 2010

Hygaboo Andersen said: Gee, it's hard to think of a more rancid combination of arrogance and greed than the evolutionists when it comes to their pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature. Not only do they charge megabucks for 24-hour access to their articles, they also demand you register with them for the privilege of sending them this money! Evolutionists are worse than the pornographers their faith spawns when it comes to their money-grubbing spam machine! Indeed, what is the point in citing this crud knowing Christians will be unable to read and critique it--or is that the whole point?! By the way, computer models of animals morphing into each other is not proof that it happened in real life! That is all this "research" really seems to amount to.

Right. Look dude, if you don't read the scientific literature, exactly why do you think that you are in any position to criticize it? If you have any evidence that anything in the scientific literature is incorrect, inaccurate, or even just incomplete, you are free to publish that evidence. If not, then kindly take your uninformed opinions and your whining and go away. By the way, most scientific journals are freely available at university libraries. Many are freely available on the internet as well. Also, "evilutionists" are not the greedy ones. They often have to pay page charges to get their articles published and then the publishers get all of the profits and royalties from the publication. If you have any beef at all, it is with the publishers, not the scientists. The point is that if "christians" are unable or unwilling to read the scientific literature, they have no one to blame but themselves. You can choose to remain in ignorance, or you can choose to become educated. What you can't do is blame someone else for your poor choices.

DS · 31 July 2010

Just to be clear, the "megabucks" charge for access to this article is $15.00 US. You could copy it at the library for about 10 cents per page. Or you could get a yearly subscription to the journal and get access to hundreds of articles for only a few dollars each.

For $15 you could probably get some snake oil from the local "christian" faith healer. Yea, that scientific literature, what a scam!

Hygaboo Andersen · 31 July 2010

John Kwok said: Nick, With the latest digitizing equipment, it's really a relative piece of cake to collect morphometric data. But people were able to use calipers and, if I'm not mistaken, Stephen Jay Gould had published extensively in the mid to late 1960s and early 1970s using a lot of morphometric data derived from calipers. Ask Kevin Padian since I am sure he might steer to some useful histories on morphometric analysis. Best, John
Nick (Matzke) said: John Kwok -- thanks, you're right I imagine, I haven't studied the history of this. However I doubt that anyone ever calipered all of the landmark-to-landmark distances, with 40 landmarks this would be 40^2 measurements per specimen. Easy with the computer, impossible without. Cheers! Nick

John, This discussion of calipers and cainine anatomy has brought me to the conclusion that since your fallout with PZ, you have taken an unholy, inordinate interest in dogs. You don't need to be ashamed since this procivity is indulged in by many evolutionists, expecially those who refer to themselves as "straight" or "heterosexual." Could you perhaps share with me the details of you have been doing with these creatures so I will be better able to help you overcome your sin. Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!

MrG · 31 July 2010

Hygaboo Andersen said: Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!

Oh, you're sleazy HA. Just gotta focus on the soft targets.

fnxtr · 31 July 2010

Wheels said: Dammit Nick, now I'm arguing with Cornelius Hunter again. Why oh why couldn't it have been Yukon Cornelius instead?

Or Jerry Cornelius. So, did dogs have more potential for variation than cats to begin with? If so, why? Or has the human selection just been more (heh heh) dogged -- focused and extensive -- in the one case?

Frank J · 31 July 2010

Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!
— HA

I suspect that the only "love and healing" you'll get is an eternity of Freshwateresque tesla coil "X-ing." You might want to practice saying "Thank you sir, may I have another?"

DS · 31 July 2010

Hygaboo Andersen said: Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!

Remember Hygaboo, no matter what you do, the article is always available to you for just $15.

Stanton · 31 July 2010

fnxtr said:
Wheels said: Dammit Nick, now I'm arguing with Cornelius Hunter again. Why oh why couldn't it have been Yukon Cornelius instead?
Or Jerry Cornelius. So, did dogs have more potential for variation than cats to begin with? If so, why? Or has the human selection just been more (heh heh) dogged -- focused and extensive -- in the one case?

Dogs have a much larger genome than cats, allowing for literally more genetic diversity, and dogs are not obligate or hypercarnivores like cats.

John Kwok · 31 July 2010

I'd love to introduce you to a hungry eight meter-long Burmese python. Would solve immediately your penchant for risible verbal diarhhea:

Hygaboo Andersen said:
John Kwok said: Nick, With the latest digitizing equipment, it's really a relative piece of cake to collect morphometric data. But people were able to use calipers and, if I'm not mistaken, Stephen Jay Gould had published extensively in the mid to late 1960s and early 1970s using a lot of morphometric data derived from calipers. Ask Kevin Padian since I am sure he might steer to some useful histories on morphometric analysis. Best, John
Nick (Matzke) said: John Kwok -- thanks, you're right I imagine, I haven't studied the history of this. However I doubt that anyone ever calipered all of the landmark-to-landmark distances, with 40 landmarks this would be 40^2 measurements per specimen. Easy with the computer, impossible without. Cheers! Nick

John, This discussion of calipers and cainine anatomy has brought me to the conclusion that since your fallout with PZ, you have taken an unholy, inordinate interest in dogs. You don't need to be ashamed since this procivity is indulged in by many evolutionists, expecially those who refer to themselves as "straight" or "heterosexual." Could you perhaps share with me the details of you have been doing with these creatures so I will be better able to help you overcome your sin. Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!

John Kwok · 31 July 2010

You have no idea what I have an inordinate fondness of.... but, wait, I think I just told you. A ravenous Burmese python slowly devouring a pathetic, intellectually-challenged Xian fool like yourself. I'd taped it and send it off to Ken Ham, Bill Dembski and their compadres as a warning as to what might transpire with them:

Hygaboo Andersen said:
John Kwok said: Nick, With the latest digitizing equipment, it's really a relative piece of cake to collect morphometric data. But people were able to use calipers and, if I'm not mistaken, Stephen Jay Gould had published extensively in the mid to late 1960s and early 1970s using a lot of morphometric data derived from calipers. Ask Kevin Padian since I am sure he might steer to some useful histories on morphometric analysis. Best, John
Nick (Matzke) said: John Kwok -- thanks, you're right I imagine, I haven't studied the history of this. However I doubt that anyone ever calipered all of the landmark-to-landmark distances, with 40 landmarks this would be 40^2 measurements per specimen. Easy with the computer, impossible without. Cheers! Nick

John, This discussion of calipers and cainine anatomy has brought me to the conclusion that since your fallout with PZ, you have taken an unholy, inordinate interest in dogs. You don't need to be ashamed since this procivity is indulged in by many evolutionists, expecially those who refer to themselves as "straight" or "heterosexual." Could you perhaps share with me the details of you have been doing with these creatures so I will be better able to help you overcome your sin. Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!

JGB · 31 July 2010

However the understanding is priceless.

DS said:
Hygaboo Andersen said: Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!
Remember Hygaboo, no matter what you do, the article is always available to you for just $15.

John Kwok · 31 July 2010

Still doesn't get it that I accept Lucifer, the Lord of Light, as my savior:

JGB said: However the understanding is priceless.
DS said:
Hygaboo Andersen said: Remember John, no matter what you do the love and healing of Christ Jesus is always available to you!
Remember Hygaboo, no matter what you do, the article is always available to you for just $15.

Frank J · 1 August 2010

However the understanding is priceless.
— JGB

For all we know he/she paid the $15, understands it, and agrees with it. What would be priceless is if we would keep the "feeding" to a minimum - just enough to show readers how selective they are with answering questions.

MrG · 1 August 2010

Frank J said: For all we know he/she paid the $15, understands it, and agrees with it. What would be priceless is if we would keep the "feeding" to a minimum - just enough to show readers how selective they are with answering questions.

HA makes a specific effort to target "readers" who are, shall we say, easily excited and show no evidence of having any ability to be selective. I do not at all believe HA is serious -- the cartoon he referenced was so over-the-top that only Ray Martinez could take it seriously. HA may not like EVIL-utionists but he's only here for the fun of it.

eric · 1 August 2010

Stanton said:
fnxtr said: So, did dogs have more potential for variation than cats to begin with? If so, why?
Dogs have a much larger genome than cats, allowing for literally more genetic diversity, and dogs are not obligate or hypercarnivores like cats.

Puny human! Believe that if you wish. We cats know that the reason is because we've reached the pinnacle of local fitness and there's no room for improvement on perfection. With dogs, well....lets just say there's lots of fitness lanscape with higher peaks. Now, be a good human and fetch me some milk and treats. [/cat] :)

raven · 1 August 2010

Getting back to the subject.

Evolution is RM + NS.

It has always seemed to me that what is rate limiting in evolution is natural selection or selection pressure.

For most of the time, species and populations seem to sit on top of local fitness optima. Not that they aren't evolving, but much of it is cryptic neutral drift.

The dog example indicates this. So does corn, maize, which is recently descended from teosinte. And of course, the many examples of adaptive radiations, Hawaiian fruit flies, Galapagos birds, African cichlids, the mammalian takeover after Chicxulub etc..

Robin · 2 August 2010

raven said: Getting back to the subject. Evolution is RM + NS.

Seems to me that based on this research (along with several others) it might be time to change the above to evolution is RM + S (or SP) where 's' is just selection or selective pressure. "Natural" selection, at this point, seems unnecessarily limiting.

Henry J · 2 August 2010

The phrase "differential reproductive success" comes to mind as a good way of stating the basic principle.

Wheels · 2 August 2010

Well, I left this comment at Hunter's blog almost two days ago. I'm waiting with bated breath for the response. Let's see what happens when Hunter isn't allowed to use his wonky Creationist definition of evolution. Can he admit to evolution being an observed fact if he can't dismiss actual evolution to talk about his Straw Man? It's kinda funny when he explicitly dismisses bacteria gaining antibiotic resistance from being an example of evolution. He has to literally define away all the undeniable examples.

John Kwok · 2 August 2010

No, Natural Selection by itself is not necessarily limiting. Anyway, Darwin used artificial selection of dogs as an analogy to explain Natural Selection in his "On the Origin of Species". But it in of itself may be incomplete in explaining such phenomena like long-term morphological stasis, which is why some biologists believe an "Extended Modern Synthesis" evolutionary theory is necessary:

Robin said:
raven said: Getting back to the subject. Evolution is RM + NS.
Seems to me that based on this research (along with several others) it might be time to change the above to evolution is RM + S (or SP) where 's' is just selection or selective pressure. "Natural" selection, at this point, seems unnecessarily limiting.

TR Gregory · 3 August 2010

Nick (Matzke) said: Hi Ryan! Thanks for the comment, I was attempting to be provocative (obviously). So I'll continue: 1. Who says these aren't permanent changes? And, who says that the morphological changes that *do* occur in "regular large scale natural evolution" *are* permanent? In either case what happens morphologically probably depends substantially on the future selective regime, which will either maintain the changes, or it won't. 2. Re: oxymoron. So, hypothetically, if a major developmental change leading to a "major" morphological change (say, on the level of the differences we typically see between genera or between families) did occur *within* a species, complete with polymorophism within the population for this change which is later fixed by selection -- but with no speciation/lineage-splitting event, would this be: (a) a microevolutionary event, because it was within a species, or (b) a macroevolutionary event, because the amount of change was on the level of the differences often observed between higher taxa? Obviously there must be a simple, correct answer, right? ;-)

Hi Nick, I forgot to come back and check the discussion. Short answers: 1. We know it's not permanent because dog populations that are not bred selectively revert to mutthood very quickly. 2. An oxymoron because the original and still current definition of "macroevolution" is evolution above the species level. Do you know of any such examples of the case you describe? If not, this seems like a minimally relevant scenario to contemplate. ps: Who the heck is this Dave C guy who thinks I might be a creationist?!? :-)

Jedidiah Palosaari · 3 August 2010

eric said: Okay, second: "by fossil evidence alone" is rarely the case, and even when it is, biologists consider more than just skull morphology. There are many other ways, for instance, to tell a wolf from a dog which make it quite clear which species is which.

The problem I see there is that, as Gould pointed out, often paleontology is how some teeth mated with other teeth to produce more teeth. Yes, we can get a hell of a lot from teeth. But not as much as if we have the entire skeleton. If our fossils are merely teeth, as they often are, it seems we, at times, would run the risk of conflating species and genera.

Jedidiah Palosaari · 3 August 2010

Hygaboo Andersen said: Gee, it's hard to think of a more rancid combination of arrogance and greed than the evolutionists when it comes to their pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature. Not only do they charge megabucks for 24-hour access to their articles, they also demand you register with them for the privilege of sending them this money! Evolutionists are worse than the pornographers their faith spawns when it comes to their money-grubbing spam machine! Indeed, what is the point in citing this crud knowing Christians will be unable to read and critique it--or is that the whole point?! By the way, computer models of animals morphing into each other is not proof that it happened in real life! That is all this "research" really seems to amount to.

I think you are unaware of the nature of science. No, that's not right. You seem to be unaware of basic English definitions. Typically, scientists *do* peer review other scientists. That's what "peer review" means. That's what science is. Unless you want to just redefine all of science? Ohhh! Right. My bad. But I see you are very anti-capitalism and free market. That of course is your right. I mean, nothing against socialism. Not that there's anything wrong with that... But you may also be confused on how to use a credit card and move your finger on the mouse pad to reach over and click. Most of the other Christians get that, I know. You may be just an incredibly stupid Christian.

Hygaboo Andersen · 14 September 2010

Jedidiah Palosaari said:
Hygaboo Andersen said: Gee, it's hard to think of a more rancid combination of arrogance and greed than the evolutionists when it comes to their pathetic, third-rate, peer-reviewed by other evolutionists, so-called literature. Not only do they charge megabucks for 24-hour access to their articles, they also demand you register with them for the privilege of sending them this money! Evolutionists are worse than the pornographers their faith spawns when it comes to their money-grubbing spam machine! Indeed, what is the point in citing this crud knowing Christians will be unable to read and critique it--or is that the whole point?! By the way, computer models of animals morphing into each other is not proof that it happened in real life! That is all this "research" really seems to amount to.
I think you are unaware of the nature of science. No, that's not right. You seem to be unaware of basic English definitions. Typically, scientists *do* peer review other scientists. That's what "peer review" means. That's what science is. Unless you want to just redefine all of science? Ohhh! Right. My bad. But I see you are very anti-capitalism and free market. That of course is your right. I mean, nothing against socialism. Not that there's anything wrong with that...

Meh, All secular political ideologies are nothing but Darwin's dingleberries. Justice can only be achieved by placing the Gospel at the center of all political discourse.

But you may also be confused on how to use a credit card and move your finger on the mouse pad to reach over and click. Most of the other Christians get that, I know. You may be just an incredibly stupid Christian.

What I am not confused about is having to sign up for spam. I don't want the moral excrement of evolutionism pollluting my email box!

Stanton · 14 September 2010

Hygaboo Andersen said: Meh, All secular political ideologies are nothing but Darwin's dingleberries. Justice can only be achieved by placing the Gospel at the center of all political discourse.

Then I take it you would prefer to live in a country like Afghanistan, Iran or Somalia, where you can be beaten or put to death for not being Godly enough, often without trial?

MuD PhD · 25 September 2010

All the journals are in fact available free to anyone at any academic library (usually found on college campuses) Not every library will have every journal but they are staffed by librarians who know how to get any article you need. The advent of paid online subscriptions is a very recent development( less than 10 years) Before that all article reading was done in librarys. There is no conspiracy to restrict access, you simply need to go to a good library!