Pharyngula

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Wednesday, August 17, 2005

Pleistocene Horses

Echoed on the Panda's Thumb

Weinstock et al. have published a solid and interesting paper that attempts to resolve some issues in the recent evolution of horses. It's good work, and shows how molecular analysis of fossils can complement morphological studies to give a clearer picture of organismal history. Unsurprisingly, though, creationists are already spluttering out nonsense about it. I'm going to give a quick overview of the scientific results, and then show some of the creationist babble in response (not too much, though—you'll quickly see how dishonest and evasive creationists are).

First, what the paper is not: it is not a big picture view of all of the evolutionary history of horses. If you want that, I greatly recommend Kathleen Hunt's horse evolution FAQ. This paper, like most science papers, is very tightly focused on a specific, addressable problem…in this case, resolving the phylogeny of just American horse species within the last 3 million years. What Weinstock et al. were trying to figure out is how 3 major groups of horse species were related to one another, and how they were related to Old World horses. The method they are using is to extract ancient mitochondrial DNA from bone specimens, sequence a particularly informative stretch, and use established computer techniques to fit them into a phylogenetic tree.

The three groups they are trying to resolve are:

  • The caballines, or the true horses, of which our modern Equus are representatives.
  • The stilt-legged horses, species that resemble the true horses but were more slender limbed, and resemble Asian asses, such as the onager and kiang.
  • Hippidion, a distinctive animal with a boxy skull and shorter, stouter limbs.

They have measurable, objective data to distinguish the three groups, as for example, in this plot of metatarsal dimensions.

horse evolution
Metatarsal Shape and Size in Pleistocene and Extant Equids. (A) Bivariate plot showing metatarsal shape and size in extant and extinct horses. Modern asses: kiang (E. kiang) = light blue circles; onager (E. hemionus onager) = dark blue; kulan (E. hemionus kulan) = purple. Pleistocene equids: stilt-legged from Alaska and the Yukon = black; E. lambei (Alaska) = red; stilt-legged from Natural Trap Cave (Wyoming) = yellow; caballines from Natural Trap Cave = orange; caballines from Alberta = green; H. saldiasi from southern Patagonia = grey. “A” and “B” above/beside points of caballine horses denote the phylogenetic clade to which the specimens belong. Not all of the specimens were genetically analyzed.

Prior hypotheses of the relationship of these three North American groups suggested that, because of their morphology similarity to Asian stock, the stilt-legged horses were related the Asian asses, and had migrated into the Americas by way of the Bering land bridge. Hippidion was so distinctly different from other horses that it was considered to have diverged from the equid lineage about 10 million years ago. These species were thought to have spread into South America during the Great American Biotic Interchange about 2.5 million years ago, when the Isthmus of Panama formed and allowed animals to move north and south between the continents. Two competing models of these relationships, based on morphological analyses, are shown below.

horse evolution
Diagrammatic Representation of North and South American Equid Taxonomy Superimposed on a Time Scale according to Paleontological Data. (A) represents MacFadden's view of hippidiform origins from a pliohippine, diversifying into two genera, Hippidion and Onohippidium, in North America during the Miocene. (B) shows Alberdi and Prado's view of hippidiforms as descendants of a pliohippine during the Miocene; the single genus, Hippidion, originates only after dispersal into South America (they do not recognize the genus Onohippidium as valid).

The molecular data revealed a surprise, though. The tree below was constructed by analysis of mtDNA, and it groups the animals very differently. The caballines of America and Europe form a single clade, as expected. The stilt-legged horses and Hippodion, though, cluster together; Hippodion diverged only recently, and the stilt-legged horses are more closely related to the caballines than to the Asian asses. The authors also suggest that many species within the caballines ought to be lumped together, are more likely to represent regional variants than true species.

horse evolution
Phylogeny of Recent and Pleistocene Equids The maximum likelihood tree was constructed with two fragments of the mitochondrial control region (583 bp and 133 bp in the HVR1 and HVR2 regions, respectively). Bayesian analysis produced a similar topology. The general time-reversible substitution model was used in both techniques. Black numbers above/beside nodes are posterior probabilities and bootstrap values, respectively (only values > 50% are shown). White numbers on black background are divergence times as calculated from the molecular data. Numbers/letters in bold at the beginning of each specimen's name are sample numbers or GenBank accession numbers. Labels of prehistoric specimens are followed by their age, if available, in thousands of years. The outgroup (Rhinoceros and Ceratotherium) is not shown.

It's a bit of a shakeup, and I'm sure taxonomists and phylogeneticists will be arguing about it for years. Here's their revised diagram of recent horse history:

horse evolution
Diagrammatic Representation of North and South American Equid Taxonomy Superimposed on a Time Scale according to Molecular Data. (C) represents the results of the molecular data presented in the Weinstock study; it shows Hippidion as originating during the Pliocene, c. 3-3.5 Ma ago. The stilt-legged horse is possibly a sister species of Hippidion.

Now what about those creationists? I was interested to discover a very recent response to this article, and I was curious to see how they would deal with it. The short answer: they didn't. They're quite sure it is wrong, of course, but they make no attempt at analysis—I suspect they just looked at it dull-eyed and uncomprehending, and did what all creationists do: went looking for some blanket disclaimer from a creationist authority that they could parrot. They found one at Apologetics Press, and it is amusingly bad. It's a rebuttal to an article by David Quammen in National Geographic that starts off this way:

In a section of his article that is an inexcusable gaffe on his part, and one that surely must represent a terrible embarrassment to his evolutionary colleagues, Quammen resurrected the long-dead concept of "horse evolution."

The concept of horse evolution isn't dead at all, and the Weinstock article is discussing at length and in some detail the evidence for evolution of several lineages. Where do creationists get this odd idea that evolutionary biologists would be embarrassed by someone discussing the evolution of horses?

It's all stone-cold ignorance, of course. The explanation comes a little further down.

Evolutionists themselves long ago abandoned horse evolution as an example of transitional forms, since they no longer believe the fossil record represents anything like a straightforward progression, but instead a bush with many varying branches.

How silly. The fossil history of horses is full of transitional forms. We don't think of evolution as simple linear progress, that is true…but even Darwin in the Origin drew a general picture of evolution as a branching tree. The idea of a linear progression is a common creationist misconception. What this article is basically doing, then, is complaining that evolutionary biologists' idea of horse history accurately reflects its complexity, while the creationists idea of simple lineages has been shown to be wrong…and they somehow mentally translate this into an example of biologists being wrong.

I really didn't need to read further. It's an excellent example of empty creationist noise against a rich (and growing richer) set of evolutionary data.


Weinstock J, Willerslev E, Sher A, Tong W, Ho SYW, et al. (2005) Evolution, systematics, and phylogeography of Pleistocene horses in the new world: A molecular perspective. PLoS Biol 3(8): e241.


Trackback url: http://pharyngula.org/index/trackback/2751/OsoZifTp/

Comments:
#36162: — 08/17  at  01:28 PM
Yay. I love it.



#36163: coturnix — 08/17  at  01:42 PM
Thanks. Now I need to re-learn everything I used to think I know about horse evolution!



#36167: Jim Anderson — 08/17  at  02:05 PM
"...I used to think I know..."

Wow. An epistemological triple-whammy.



#36170: coturnix — 08/17  at  02:16 PM
I LOVE epistomological triple-whammies! wink



#36171: — 08/17  at  02:18 PM
An absolutely beautiful quote from that creationist site:

Interestingly, another editor of a well-known magazine tried this tact several years earlier, and ended up being publicly scolded for it. John Rennie, editor of Scientific American, wrote in the July 2002 issue of that publication: “Actually, paleontologists know of many detailed examples of fossils intermediate in form between various taxonomic groups…. A sequence of fossils spans the evolution of modern horses from the tiny Eohippus” (2002, 287[1]:83).

"AHA! Yeah, take that, you evilutionists! Even your own kind admit... that... there... is... a... sequence of transitional forms.

"Um.

"Can I get back to you on this?"

Absolutely amazing. I'm trying to think of ways that the Rennie quote can possibly be any comfort to the creationists, but I can't come up with a single one. It's like they can't even comprehend simple, clear English, let alone the technical language of science. They just look for signs of debate within the scientific community, and have arbitrarily decided that one side must be on the side of 'evilution', and the other side, creationism. Absolutely stunningly stupid.



's avatar #36175: PZ Myers — 08/17  at  02:34 PM
I think the creationist saw Rennie making a clear statement about the evolution of horses, and then since he has convinced himself that biologists have abandoned the notion of horses evolving, he imagined all them ding-danged evilutionists rebuking him.

Remember, these people hear voices in their heads and think it's god having a chat with them.

PZ Myers
Division of Science and Math
University of Minnesota, Morris



#36177: paperwight — 08/17  at  02:41 PM
One might also think that if they're going to be creationists, they should at least be grammatical creationists. There's no such thing as "trying a tact". Itwould be "trying a tack", or more often, "taking a different tack", based on the nautical term "tack".

I can't take these people seriously anyway, but when they can't even be bothered with proper grammar, it tends to underline their incompetence.



#36185: Jim Harrison — 08/17  at  03:31 PM
Every statement, true or false, material implies every true statement. So if you are absolutely, positively sure of something, you know in advance that everything you read or hear implies it. For example, since Creationists just know that they are right, they rightly assume that every new piece of research supports their views. Of course, material implication is like kissing your aunt; but, as Humpty Dumpty would triumphantly conclude, "It's logic!"



#36189: Kele — 08/17  at  04:13 PM
Heh, my dad and I recently got in a little conversation about horse evolution in the comments part of one of my blog posts. He's on the evolution side but he doesn't really know much about it so he offered me links that said horse evolution was wrong and it's presented in a wrong way in textbooks and stuff like that. It's great timing that you posted about this because now I can show him horse evolution is still alive and well. Very interesting, by the way.



Trackback: Speaking of horse evolution... Tracked on: Kele's Atheistic and Evolutionary Journey (72.9.234.70) at 2005 08 17 16:15:09
PZ Myers discusses a recent paper on horse evolution. See? Horse evolution hasn't been refuted and it's still very much alive. (I think this mainly goes to my dad, haha.)



's avatar #36194: Chris Clarke — 08/17  at  04:44 PM
I'm sorry, I don't see any mention of fox terriers here. Isn't that against the law or something?

"I do not think we should antagonize the religious when it is not warranted, though I think we should be willing to do so whenever it is.”
-- Glen Davidson



#36196: — 08/17  at  04:53 PM
Has anyone studied the evolutionary track from the rather small rear end of eohippus to the modern day horse's asses at the Discovery Institute?

-jcr



#36198: — 08/17  at  05:05 PM
What will the Braminologists do? They have analyzed the equids http://www.grisda.org/origins/57009.pdf using a multi dimensional scaling method, similar to XGobi http://www.research.att.com/areas/stat/xgobi/
They don’t like molecular data. It doesn’t work with their systematic methods i.e. based on primates, it doesn’t yield the correct results.
http://www.creationresearch.org/crsq/abstracts/sum34_4.html

If the proposed relationship between stilt-legged horses and Hippodion and other changes are supported by additional data the Baraminologists are not only stuck with fitting everything into 6000 years but explaining why molecular data is unacceptable.



#36212: — 08/17  at  06:08 PM
Remember, these people hear voices in their heads and think it's god having a chat with them.

I think there's a very interesting connection between delusions seen within creationism and those seen within schizophenia. In the latter it's notable that intricate sustained delusional systems are mostly seen in highly intelligent sufferers, because the less intelligent are incapable of integrating their bizarre intuitions into a pseudo-coherent whole. Consequently, more intelligent psychotics are often harder to treat because they have semantic structures which are resistant to unlearning even after they are biologically responding to treatment. It seems like people who are desperate to integrate irrational facts get caught up in very similar cognitive processes, even though the underlying reasons are non-biological.



#36215: — 08/17  at  06:47 PM
PZ: Ah, that makes slightly more sense then. What threw me was the actual lack of any quotes from scientists "publicly scold[ing]" him, so I thought they meant Rennie's quote was the "scolding."

Perhaps, then, the author was referring to the incoherent screams of denial emanating from his own camp. Creationists are so cute when they think they're contributing to the scientific dialogue.



#36224: Jeff Fecke — 08/17  at  07:47 PM
If horse evolution is true, then how do you explain GIANTS AND DWARVES?

Ha! Now you're in a box!



#36227: — 08/17  at  07:58 PM
I suppose IDists may say "OK horses are not designed, but how about those unicorns!"



#36229: coturnix — 08/17  at  08:06 PM
How about Falabellas and Shires? (Better than the fox terrier, anyway).



#36238: — 08/17  at  08:58 PM
Hey Jeff... LOL
I forgot that one. It was so abused for a week... Now it is a thing of beauty.



#36242: Joseph ODonnell — 08/17  at  09:18 PM
It can never truly die MpM.



#36250: — 08/17  at  10:47 PM
Lest anyone be misled by the lack of clarity in the genetic portion of the analysis - no DNA older than 53,000 years was studied and it was all from a single species of equus.

http://biology.plosjournals.org/archive/1545-7885/3/8/supinfo/10.1371_journal.pbio.0030241.st002.doc



#36253: — 08/17  at  11:44 PM
"OK horses are not designed[...]"

"How about Falabellas and Shires?"

"Proof that you can detect design!"

If an animal "fits" its environment, it was obviously designed to be there. Since evolution doesn't occur. Because there was a designer. Because evolution doesn't occur.



#36254: coturnix — 08/18  at  12:00 AM
If an animal "fits" its environment, it has obviously evolved there. Since evolution is the process that produces adaptation. Because there is no designer. Because evolution occurs.

Convinced yet?

Falabellas and Shires are results of artifical selection.

Were horses designed for riding or for pulling a cart?

Wish we could discuss fine points of niche-construction, hierarchical units of selection, or developmental constraints instead of repeating the basics all over again....



#36264: — 08/18  at  05:18 AM
Lest anyone be misled by the lack of clarity in the genetic portion of the analysis - no DNA older than 53,000 years was studied and it was all from a single species of equus.


First off, congratulations on ferreting out the fact that all the mtDNA samples were all less than 53,000 years old - a statement diabolically concealed right in the first paragraph of the Results/Discussion section. Leaving aside the fact that this is a surprise to no one who's seen mtDNA studies before, if it weren't for your vigilance, lay readers may well have thought that the devious scientists were out to claim that their results were representative of horses who lived ca. 800 billion years ago.

Second, I'm not entirely clear on how it is that you can claim all samples came from a single species of Equus. Looking at the list you linked to, it says that samples 1-33 and 42-53 are attributed to Equus sp. Samples 34 and 35 are designated Equus cf. kiang. Samples 36-40 represent Hippidion saldiasi and sample 41 is Hippidion principale.

Obviously, none of the Hippidion samples represent "a single species of Equus" as they come from an entirely different genus. Moreover, the "sp." in "Equus sp." means that no species is specified or determined, meaning a whole slew of species may be represented in the sample.

Did you perhaps mean to say all the mtDNA examined was "from a single basic kind of horsey"?



#36269: — 08/18  at  07:52 AM
"Yeah, but it's still a horse."

Nyuk, nyuk, nyuk.



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