Phylogenomic Fallacies
This is the fourth in a series of articles for the general public focused on understanding how species are related and how genomic data is used in research. Today, we talk about some common fallacies in phylogenomics.
Where do humans fit on the evolutionary tree of life? This is an important topic in evolutionary biology. A lot of people believe humans are the most important and highly-evolved organisms, but in reality, all modern species are equally evolved. Our natural tendency to assume that humans are evolutionarily superior has led to a few misconceptions about phylogenetic trees.
To understand the first misconception, let's look at a phylogenetic tree of plants (from "The Amborella Genome and the Evolution of Flowering Plants"). Eudicots and monocots are two classes of flowering plants, or angiosperms, and the plants in black are non-flowering plants. The term "basal" refers to the base of a phylogenetic tree, and a basal group is a species that branches closer to that base. The authors chose to label the angiosperms that are not eudicots or monocots as "basal angiosperms." But this label is arbitrary; all the angiosperms are equidistant from the common ancestor and thus equally evolved. We sometimes tend to give more weight to branches that contain the species of interest and call other branches basal, almost assigning them a lesser importance. In this case, the species of interest is plants that consist of many foods that humans eat; a species is often deemed more important as it relates to humans. But modern species are equally evolved from a root common ancestor regardless of when their branch diverged from the common ancestor. To avoid confusion, it might be best to eliminate the "basal" term altogether.
This type of thinking also leads us to place humans at the end of phylogenetic trees. However, this placement is arbitrary and trees can be drawn in many equivalent ways. For example, compare a tree of primates with the branches rotated. The tree on the left, with humans at the top of the tree, is one you might see more often. But both of these trees are actually identical, and the relationships between species that can be inferred from the tree on the right is the same as the relationships in the tree on the left. Species at the tip of a tree are equidistant from the root common ancestor, so they can be considered evolutionarily equivalent.
Similarly, a common misconception is that humans evolved directly from monkeys. Monkeys, though, are modern species just like we are and have been evolving and changing over time. The common ancestor we share with monkeys may have looked much different than monkeys do now. This assumption that modern species represent an ancestral state of human evolution is what T. Ryan Gregory calls the platypus fallacy. Gregory uses the example that we can't examine the traits of platypuses and think that humans at one point in their evolution possessed these same traits. We can no more infer the traits of human ancestor species from platypuses than platypuses can infer the traits of their ancestors from us.
Human-centered thinking is very prevalent in our society, affecting our laws, religions, and customs. While it probably influences all of us on a personal level, it can lead to false conclusions and misconceptions in science, like thinking that humans are the most highly evolved species. But all modern species are evolutionarily equivalent because they have been evolving for the same amount of time. Eliminating this fallacy will enable us to better understand the evolutionary process.
For more information on basal groups, check out: "Which side of the tree is more basal?, Krell, Frank et al. Systematic Entomology (2004).
This series is supported by NSF Grant #DBI-1356548 to RA Cartwright.
136 Comments
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 16 December 2014
Well, of course humans evolved directly from monkeys. Humans are monkeys. I know the fallacy you're attacking here, but that isn't it. I'm not complaining about the principle, in other words, just the specific example you choose. In fact, the common ancestor of humans, guenons, and tamarins would presumably have possessed whatever characteristics are shared between guenons and tamarins. We can't pick out one species and say that it's like the ancestor, but we can examine how traits are shared over the tree. The problem here is that "monkey" as commonly used refers to a paraphyletic group in which humans are embedded.
DS · 16 December 2014
Excellent post. This misconception goes all the way back to Aristotle and the Scala Naturae. Humans just naturally think in hierarchies. However, this can be a very misleading concept in phylogenetics. And creationists aren't the only ones who get this wrong. Showing the two trees side by side is an excellent way to get the point across. As for the term basal, it does have a specific meaning, even if it is often used incorrectly or interpreted incorrectly. At least the terms basal and derived carry less negative connotations that the terms primitive and advanced.
Creationists usually claim that humans evolved from chimps. This is of course incorrect, as can clearly be seen in either version of the tree. However, to claim that humans evolved from monkeys might be correct, depending on your definition of that term, as correctly pointed out by the previous poster. Perhaps it would just be best to say that humans evolved from other primates.
Palaeonictis · 16 December 2014
harold · 16 December 2014
The entire biosphere is all the product of billions of years of evolution since the origin of cellular life, so the term "more evolved" is meaningless.
In fairness, some anatomic, physiologic, or molecular features are more recent that others. Cell membranes and the citric acid cycle are extremely ancient features of life, for example, whereas internal skeletons in animals is a much more recent feature.
Humans do have some unique recent features, such as our particular bipedal gait (bipedalism itself is fairly ancient but not our type), or particular brain structure, our speech (recent evidence shows that bird speech, which is far more ancient than human speech, may be surprisingly related, but our speech evolved independently from theirs). It's mildly interesting that birds have a form of speech, bipedalism, and specialized forelimbs, just like us.
So humans are clearly not "more evolved" than anything else in the biosphere, but we do have many recent traits.
It's often conjectured that some types of organisms like amoebae have relatively fewer recent traits and have been more similar to distant past ancestors for longer. This is hard to confirm but may not be totally unreasonable.
eric · 16 December 2014
harold · 16 December 2014
https://me.yahoo.com/a/JxVN0eQFqtmgoY7wC1cZM44ET_iAanxHQmLgYgX_Zhn8#57cad · 16 December 2014
Yes, I was thinking that when you can recognize a coelacanth in a fish market when only fossil coelacanths were known, coelacanths are really a bit closer, at least in morphology, to ancestral lobe-finned fishes than is a gorilla. Mosses, liverworts, while certainly not unchanged in morphology, remain recognizable, while angiosperms have recognizably changed more. And adaptive radiations seem to be real--Cambrian and all of that.
Rates of evolution aren't all that clear. Is genetic change really a good way to date splits, or do some lineages accumulate change more slowly. But then there's the question of what kind of change is going on. Possibly coelacanths have changed genetically every bit as much as our lineage has, but if coelacanth changes involved largely neutral changes, is that what we mean by "evolved a lot"?
Then there are generational times that might affect evolutionary rates. Mice presumably could evolve much faster than humans, although in morphology our line seems to have changed more, at least recently. Do bacteria and archea that live and reproduce very slowly evolve as fast as, say, E. coli?
It does seem that evolutionary "speeds" are likely to differ, certainly by some measures, such as morphology. If this were not so, how would life bounce back after major extinctions?
Glen Davidson
https://me.yahoo.com/a/JxVN0eQFqtmgoY7wC1cZM44ET_iAanxHQmLgYgX_Zhn8#57cad · 16 December 2014
Of course, the "most evolved" genomes belong to viruses.
Glen Davidson
DS · 16 December 2014
Interesting that whole genome duplication events occurred in two of the most successful lineages, angiosperms (red star) and vertebrates (which had two).
eric · 16 December 2014
harold · 16 December 2014
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 16 December 2014
DS · 16 December 2014
Well other than insects, these are the two most diverse and dominant groups in the terrestrial environment. They have certainly been highly successful in terms of speciation and diversification. Ferns cannot have more common genome duplications, since all angiosperms and vertebrates posses them as a derived character. This is evolutionary polyploidy rather than regular polyploidy. It involves not just duplicate numbers of chromosomes but significant neofunctionalization and diploidization. Of course ferns are also successful in the sense that they still exist and continue to evolve. But they are certainly not as successful as angiosperms or vertebrates by most reasonable measures. My apologies to fern lovers everywhere.
The point is that gene duplication, sometimes through whole genome duplication, provides the raw material for mutation and selection, sometimes making entire lineages highly successful. So Ohno was right when he pointed this out 45 years ago.
Robert Byers · 16 December 2014
Do humans fit on a tree? Is there any bio sci evidence to justify this tree concept beyond mere looks/dna-looks??
People think we are more evolved because we are smarter. Its not about feet.
If evolution is about changes then why not the more changes there are the more one has evolved? not better but more!
is the author against human centered concepts in civilization? just a joke!
NO EVIDENCE , again, was presented from bio sci to demonstrate to a critic, a slight majority etc in North America, as to why peopler/plants are evolved from common descent place.
Its just presumed. Looks is not bio sci evidence.
A creator would also do it this way. Basic plans would easily explain like looks at basic levels.
Man/ape being a special case because we are special. Thats why only us study these things.
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 16 December 2014
DS · 16 December 2014
Robert Byers · 16 December 2014
DS · 16 December 2014
Mark Sturtevant · 16 December 2014
I think that statements like saying 'humans evolved from monkeys' or that 'humans evolved from apes' ignores the detail that the terms 'monkey' and 'ape' is meant to refer to modern species. If one wants to be strictly formal, it is best to say that 'humans and monkeys (or apes) evolved from common ancestors'. For example, the great apes are modern primates that walk around on their knuckles but our lineage likely never did that.
Similarly, to say that 'humans are monkeys (or apes) runs into a similar problem. I know that such constructions are desirable for cladistics, but I think the phrase to use here is that 'humans and apes are both hominids'.
As for the basal group fallacy, I agree entirely with the descriptions above. But I hold out that it is ok to refer the the characters that are prominent in a group to be basal. So the characters of a lung fish are basal to the lungfishes and tetrapods.
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 16 December 2014
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 16 December 2014
https://me.yahoo.com/a/JxVN0eQFqtmgoY7wC1cZM44ET_iAanxHQmLgYgX_Zhn8#57cad · 16 December 2014
KlausH · 16 December 2014
The "platypus fallacy" comment makes little sense to me. If a human and a platypus are closely examined, it would be obvious that they were related, and it would be easy to deduce some traits of the common ancestor, such as internal skeleton, DNA, four limbs, paired eyes and lungs, et cetera. One would also get a good idea as to how and when the derived characteristics appeared.
DS · 16 December 2014
4ras wrote
"Again, my claim: there is no particular relationship between whole-genome duplication and diversity."
Again, there is good evidence that this is not true in vertebrates and probably not true in angiosperms. There are also good reasons to suppose that it might be generally true. I suppose the only way to tell if this is a statistically significant relationship is to do a statistical analysis. A few examples and a few counter examples are not sufficient to resolve the matter conclusively.
DS · 16 December 2014
Here is a recent review article that seems to indicate that there are still many unanswered questions in the area under discussion:
Madlung (2013) Polyploidy and its effect on evolutionary success: old questions revisited with new tools. Heredity (2013) 110, 99â104
robert van bakel · 16 December 2014
Well done! After an early scare, ignoring Byers tantrum has worked, and I can read in peace.
Yardbird · 16 December 2014
Emily Thompson · 16 December 2014
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 17 December 2014
https://www.google.com/accounts/o8/id?id=AItOawkbohD6goiMSfag_XaXq20elmGG7Gs4tas · 17 December 2014
Mike Elzinga · 17 December 2014
Thinking back to Andreas Wagner's recent book, The Arrival of the Fittest, and his discussion of "genotype networks," I suppose this concept is more applicable to large molecules in which a network of genotypes produce essentially the same phenotype; say a number of different genotype molecules that perform essentially the same metabolic processes.
I understand that there are some precedents for such an idea; some of it coming from the fact that, in many organic compounds, it is the shape of the molecule and not necessarily the specific atoms at given locations that determine the reactive properties of the molecule.
It prompts the question about what happens when many layers of complexity are added on top of this as an organism becomes more complex; namely, can the same genotype produce different phenotypes that are reproductively isolated? Alternatively, could different genotypes produce essentially the same phenotype at these higher levels of complexity? I presume the latter case would be related to "convergent evolution."
I guess the more general question would be, "Where along the scale of complexity does speciation occur?" Is is necessarily at the level of the DNA involved in reproduction?
More specifically, if the molecular processes become more susceptible to ambient physical conditions as increasingly complex molecular assemblies start interacting with smaller binding energies (e.g., we already know that temperature during incubation can determine sex in some animals), couldn't mutations and variations lead to sufficiently different phenotypes that speciation can occur above the level of the DNA involved in sexual reproduction? Wouldn't that be genetic isolation?
What is evo-devo telling us?
Mark Sturtevant · 17 December 2014
TomS · 17 December 2014
I think that one of the problems about the names for the primates is specific to English: We have the two words, "ape" and "monkey". "Ape" is the older term, and, before Europeans started exploring, the only non-human primates known were what we now call "Old World monkeys" - that is what were called "apes". For example, that is what the King James Version means by "ape". Somebody decided to change the designation after the other primates were discovered and the word "monkey" was introduced to English. It could have been that new word "monkey" was restricted to New World monkeys, and "ape" would have been kept for Old World monkeys and the newly discovered gorillas, etc.
Palaeonictis · 17 December 2014
DS · 17 December 2014
harold · 17 December 2014
DS · 17 December 2014
From the abstract of the paper that demonstrated the two vertebrate duplication events:
"Our results highlight the potential for these large-scale genomic events to have driven the evolutionary success of the vertebrate lineage."
PLOS Biology (2005) 3(10):e314
So it seems that, at least in this case, the duplications were essential to the success of the lineage.
DS · 17 December 2014
harold · 17 December 2014
less evolved, just kidding, have more conserved ancient traits, such as fish and salamanders. So a tetraploid ancestor is hardly a shocking idea.John Harshman · 17 December 2014
DS · 17 December 2014
OK. Then here is a more direct test off exactly how whole genome duplication has led directly to diversification:
Hoffman et. al. (2012) WHole-genome duplication spurred the functional diversification of the globin gene superfamily in vertebrates. Molecular Biology and Evolution 29(1):404-423.
From the abstract:
"The physiological division of labor between the oxygen transport function of hemoglobin and the oxygen storage function of myoglobin played a pivotal role in the evolution of aerobic energy metabolism, supporting the hypothesis that WGDs helped fuel key innovations in vertebrate evolution."
Now of course this doesn't prove that whole genome duplications always lead to diversification, or that they are required for diversification, It only proves that they can, and for good reason. It would be premature to declare that they don't have anything to do with evolutionary success, since this is a direct test and confirmation of exactly that hypothesis.
This discussion is getting off topic. I will respond further on the bathroom wall if anyone wants to continue.
mattdance18 · 17 December 2014
John Harshman · 17 December 2014
DS · 17 December 2014
Agreed.
harold · 17 December 2014
Well, it's a relief to know that cockroaches have only been plaguing the Earth since the Cretaceous, or at worst, late Jurassic period. Unfortunately that does still make them a persistently successful phenotype that has survived major extinction events.
Just Bob · 17 December 2014
Henry J · 17 December 2014
Henry J · 17 December 2014
I've noticed that the http://tolweb.org/tree site appears to avoid using clade names that have colloquial meanings different than the evolutionary meaning. That makes sense to me; is it a widespread practice in the field?
John Harshman · 17 December 2014
Henry J · 17 December 2014
Specific?
Eukaryotes
Opisthokont
Metazoa
Bilateria
Deuterostomia
Chordata
Craniata
Vertebrat
Gnathostomata
Teleostomi
Osteichthyes
Sarcopterygii
Terrestrial Vertebrates
Tetrapod
Anthracosauria
Amniota
Reptilia
Synapsida
Eupelycosauria
Sphenacodontia
Sphenacodontoidea
Therapsida
Theriodontia
Cynodontia
Mammalia
Eutheria
Primates
Hominidae
Most of those are (AFAIK) technical terms that don't have common usage meanings that are paraphyletic groups. Not quite all of them, though.
(I didn't want to double space that list, but I don't know how to get it to insert a line break without a blank line between items.)
Henry
John Harshman · 17 December 2014
Joe Felsenstein · 18 December 2014
In reply to Mark Sturtevant and John Harshman:
Another way of addressing the issue of whether we are descended from monkeys is this: If ceboids (New World monkeys) and cercopithecoids (New World monkeys) are both to be called monkeys, what about their common ancestor? If we saw it, would be call it a monkey? It certainly wouldn't look like a hippopotamus! I am pretty sure that we would also call it a monkey. If a layperson were to see it, and we didn't call it a monkey, they would ask why.
Of course, it would be our ancestor too, and an ancestor of all the apes. So yes, we are descended from a "monkey".
Similarly if we saw the common ancestor of all apes (including us), we would certainly call that an ape. It would look like an ape, and people's eyebrows would be raised if we didn't call it that. So we are descended from apes.
I'd go farther and say, even without making a technical taxonomic definition of "ape" and "monkey", that this means that we are apes and that we and all the apes are also monkeys. I can see that other people might not join me in that. They might point out that I probably wouldn't also say that we are worms, even though the common ancestor of all things called worms might look an awful lot like a worm, and it would be our ancestor as well.
harold · 18 December 2014
The common English word "monkey" does not include humans in ordinary usage, and I can prove it right here with a simple thought experiment. I hope in advance that no-one is dull enough to imagine that I am denying the close evolutionary relationship between humans and our fellow primates, just making a point about language.
I'm going to show you a picture of a spider monkey and a human. The spider monkey is holding a banana and the human is holding an orange.
Now I'm going say "Relying on conventional usage of the English language in conversation, what is the monkey holding"?
If you say "I need more information, there are two monkeys", you're being pedantic.
The word monkey is a word that is informal, but that usefully distinguishes humans (and many other things) from a different group of primate.
TomS · 18 December 2014
And let's say we sought a response with a marmoset, a baboon, a gibbon, and a tarsier. I wonder how many people would treat the marmoset and the baboon as the only two monkeys; and the gibbon as the only ape?
fnxtr · 18 December 2014
Palaeonictis · 18 December 2014
https://me.yahoo.com/a/JxVN0eQFqtmgoY7wC1cZM44ET_iAanxHQmLgYgX_Zhn8#57cad · 18 December 2014
mattdance18 · 18 December 2014
Just Bob · 18 December 2014
I think the sense of the cockroach meme is a philosophical taking-down of the 'humans rule the world' idea, with an ick factor kicker: "Cockroaches have been here since forever and survived stuff that could kill off the dinosaurs. We've only been here since yesterday. Who do you think will be here after our global thermonuclear war/asteroid strike/biowarfare total genocide?"
Reed A. Cartwright · 18 December 2014
Palaeonictis · 18 December 2014
AltairIV · 18 December 2014
Robert Byers · 18 December 2014
Palaeonictis · 18 December 2014
Scott F · 18 December 2014
TomS · 19 December 2014
Palaeonictis · 19 December 2014
Just Bob · 19 December 2014
mattdance18 · 19 December 2014
RWard · 19 December 2014
Just Bob, if Paleonictus thinks he has a chance of educating Robert Byers he's sadly mistaken. BUT his efforts are not necessarily in vain. There are those lurking on Panda that may indeed profit from his response to Mr. Byers' foolishness.
That's the value of R. Byers, FL, and their ilk. Their nonsense provides an opportunity to respond to creationist arguments. Besides, they make their own side sound ridiculous. Don't bannish them to the Wall, honor them...
Robert Byers · 20 December 2014
Robert Byers · 20 December 2014
Palaeonictis · 20 December 2014
Robert Byers · 20 December 2014
phhht · 20 December 2014
DS · 20 December 2014
Yardbird · 20 December 2014
Yardbird · 20 December 2014
Yardbird · 20 December 2014
gdavidson418 · 20 December 2014
Yardbird · 20 December 2014
John Harshman · 21 December 2014
DS · 21 December 2014
DS · 21 December 2014
The paper cited above does indicate that there are times when it is appropriate to use the term "basal". This term does have a definite meaning. However, it also states that it would be more appropriate to describe the relationship as a sister group. So in this case it would be more correct to say that marsupials are the proper sister group to the placentals. That description has the added advantage of also falsifying the hypothesis that modern placentals evolved from modern marsupials, or that they magically poofed into existence after the flood for some reason. Either way, bobby is just plain wrong again.
Palaeonictis · 21 December 2014
John Harshman · 21 December 2014
Palaeonictis · 21 December 2014
DS · 21 December 2014
mattdance18 · 21 December 2014
Byers' creationist take on marsupials. Enjoy.
http://www.nwcreation.net/articles/marsupial_migration.html
(Feel free to answer my longstanding questions about your "speculation" any time, Robert.)
Palaeonictis · 21 December 2014
mattdance18 · 21 December 2014
The irony, of course, is that Byers himself assumes that when a reproductive system is termed "primitive," this means something evaluative, and negatively so, rather than just descriptive -- thereby illustrating exactly the sort of misconception this post is supposed to dispel.
But then I wouldn't expect someone whose writing exhibits such a shaky grasp of basic grammar to exhibit a firmer grasp of ideas and reasoning.
Palaeonictis · 21 December 2014
John Harshman · 21 December 2014
Palaeonictis · 21 December 2014
Palaeonictis · 21 December 2014
Creationist interpretation of human evolution:
Since Man exists, monkeys must not exist according to evilutionists.
Scott F · 21 December 2014
DS · 21 December 2014
Scott F · 21 December 2014
John Harshman · 21 December 2014
Palaeonictis · 21 December 2014
Palaeonictis · 21 December 2014
Robert Byers · 22 December 2014
Robert Byers · 22 December 2014
Robert Byers · 22 December 2014
phhht · 22 December 2014
mattdance18 · 22 December 2014
mattdance18 · 22 December 2014
John Harshman · 22 December 2014
John Harshman · 22 December 2014
Yardbird · 22 December 2014
mattdance18 · 22 December 2014
Palaeonictis · 22 December 2014
Scott F · 22 December 2014
John Harshman · 22 December 2014
Palaeonictis · 22 December 2014
mattdance18 · 23 December 2014
Palaeonictis · 23 December 2014
John Harshman · 23 December 2014
Robert Byers · 24 December 2014
Robert Byers · 24 December 2014
phhht · 24 December 2014
Palaeonictis · 24 December 2014
John Harshman · 24 December 2014
mattdance18 · 24 December 2014
Henry J · 24 December 2014
mattdance18 · 24 December 2014
Just Bob · 24 December 2014
"And what exactly were kangaroos, when they were still on the Ark?"
Ooh! Ooh! Can I answer?
Well, they hop around on two legs... so what else does that? I Know! Birds! So kangaroos are just descendants of the Bird Kind, adapted for Australian conditions. (How'm I doin', Bobby?)
But wait, no... What animal on the Ark had a body most shaped like a kangaroo? Two big hind legs, smaller front legs, a long, heavy tail? The Tyrannosaurus rex! That's who moved to Australia and
evolvedadapted itself in a couple hundred years to become a marsupial mammal!Gee, what's so hard about this bio sci stuff? I can do it without no PhD!
Scott F · 24 December 2014
Henry J · 24 December 2014
Palaeonictis · 24 December 2014
Palaeonictis · 24 December 2014
John Harshman · 24 December 2014
Palaeonictis · 25 December 2014
Dale · 28 December 2014
Even the way we name ourselves and the groups we belong to shows our human-centered biases. Like naming ourselves "Homo sapiens" (wise man, do we really deserve to be considered wise?), Primates (prime means first, so the order we are in is the first, right?), or even the calling of the sub-phylum we are in the Vertebrates and referring to all others as "Invertebrates" as if having a backbone makes us more special than insects or mollusks.
Palaeonictis · 28 December 2014
arlin.stoltzfus · 20 January 2015
The author seems to address one fallacy by proposing another, namely that "all modern species are evolutionarily equivalent because they have been evolving for the same amount of time." If she had said all modern species are equivalent *in only the sense that* they have evolved the same amount of time, that would have been defensible. The cited reference is only about the fallacy of calling one sister clade "more basal" than another, and does not justify the grand claim that there is only one factor, time, that affects evolution.
Species are NOT evolutionarily equivalent. Lineages experience different conditions. They may have different population sizes. Clades may show different rates of speciation and extinction. Prokaryotic species have evolved under high population sizes for billions of years. We have not. Species show different degrees of cohesion depending on how sexual they are. In molecular analyses, some lineages consistently have higher rates of change. They have literally evolved more.
Furthermore, these asymmetries are not independent of post-hoc tree shape recovered in molecular analyses. If divergence in characters is linked to speciation in any way, then there will be asymmetries in phylogenies. Dealing with characters that affect speciation is an actual area of research (e.g., Maddison, Midford and Otto 2007). This isn't my area so I'm not sure how broad is the literature.
I certainly would not dismiss the possibility that non-equivalent modes of evolution are linked to the asymmetry that one often sees in phylogenetic trees, in which the tree is much more ladder-like than one expects from a model assuming equivalence.