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Taxonomy questions: Aves

Badly Shaved Monkey

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Reading through randman's recent thread:

http://www.internationalskeptics.com/forums/showthread.php?t=215456

I've raised for myself questions that I've bumped into before and couldn't answer properly.

I'm confused by the taxonomy of birds. Education by Wikipedia has failed me. I think the problem is that my school biology long predated modern cladistics, so I've only learned about modern taxonomy in a rather piecemeal way.

Here,

http://en.wikipedia.org/wiki/Avialae

Aves is a subgroup a long way down the hierarchy within class Reptilia. Oddly, the page links to Aves here,

http://en.m.wikipedia.org/wiki/Aves

where Aves is a class just like I learned in school biology 30 years ago.

Am I right that these classifications have different organising principles?

Can someone point to a tree diagram that neatly lays the two approaches side by side so I can compare them?

Accepting that these classifications represent different approaches, nonetheless the gap between Aves being a Class and merely a Subgroup seems enormous. Am I right to find it strange that such a high level division by one system can be such a low level division by the other?

Obviously, the evolution of birds is particularly controversial within evolutionary theory and in confrontation with creationists, but are there other areas in taxonomy that are disputing over such a large change in hierarchical level?
 
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There's nothing strange about a group seeming higher in one system than it is in another one that works differently. It happens in every case in which large chunks of a clade are extinct and the group you're looking at is the only remnant, because one system doesn't look at extinct species and the other does.
 
Thanks for that. Is their a simple term to describe the taxonomy that has Aves as a Class to distinguish it from the cladistic one?

I think I see the point you are making except that under the old-style taxonomy we were still able to call dinosaurs Reptilia, so it can't be completely correct that no account was taken of extinct species.

Could you give an example like Aves that got nested in to the hierarchy at a much lower point using cladistics?
 
LOL, Badly Shaved Monkey, I don't blame you for the confusion ;)

Aves as a Class goes back at least to Linnaeus. It took a while (and a lot of work!) to work out the phylogentic tree as it stands now. Now, as far as I understand it, birds are still a monophyletic group, but the old "Reptilia" is not. You can keep 'Reptilia' as a Class if you include 'Aves' under 'Reptilia'... and confuse everyone who hasn't been keeping up with this stuff :P

Early classification schemes suggested that birds and "reptiles" split from each other after mammals and reptiles (or birds). Those schemes, however, also suggested reptiles all radiated from each other after the split from birds.

Relatively new information suggests instead that some "reptile" groups radiated from each other prior to the appearance of the 'Aves' group. IIRC, the first hint of this was that birds seemed more similar to crocodilians than to other reptiles; initially it seemed that perhaps an ancestral reptile similar to crocodilians would explain that (didn't seem to work, though). As more information was acquired, it seemed more and more evident that crocodilians were in fact more closely related to birds than to other 'reptiles'.

Wikipedia's entry for 'reptile' has a nice diagram showing the polyphyletic problem relatively clearly:
http://en.wikipedia.org/wiki/Reptile

Kingdoms, Classes, Orders, Species, etc. can still act as handy short-form for talking about groups of organisms, but it's becoming more and more apparent that the borders are not nearly as clean as most folks would like them to be... I'm not sure it's clear that we should keep 'Reptilia' as a Class and relegate 'Aves' to a lower subgroup, as opposed to keeping 'Aves' as a Class and reorganize 'Reptilia' to a higher-level group, or if we should just toss everything and call each level a 'subgroup', or some other option. At this point, I'm planning to let the taxonomists sort out the semantics and let me know what they decide ;) Personally, while I do need to know the relationships between and within groups, what they're actually named doesn't really matter, as long as I have something more-or-less standard to call them in a given publication...

Of course, the 'tree of life' remains a work in progress. There's no reason to believe we won't continue to modify it as we learn more, so be prepared :p

I wasn't able to find a nice (and accurate... I'm sure my recollection's a bit off!) historical account of the changing placement of 'Aves', although I confess I didn't really try all that hard. If anyone knows of one offhand, I wouldn't mind reading it... although I'm sure it's not really finished yet!
 
Thanks, I think I'm a bit further on. Yes, the Reptile page at Wikipedia helped.

Following links on from there to the Paraphyly page, I find this;

"Paraphyletic groups often have their origin in traditional taxonomy, based on similar morphological characteristics. The original perception may have been that the group was entirely descended from a single ancestor."

I think I get it a bit better now. Really, the problem is that old Reptilia was paraphyletic because the historical taxonomists saw birds as being very different from reptiles, but internally consistent. Actually the latter is true, birds are monophyletic, but the former is not, if you exclude them from reptiles then old-style reptiles turn out not to be monophyletic.

Right so far?

I need another prompt now. If Reptilia is the class containing birds, do we have a word that describes the paraphyletic group that would represent was we call reptiles in common language? Is there any use for such a word?
 
One more thing, going back to the Paraphyly page, it does actually answer my request for other examples like Reptilia/Aves. Dicotyledons, Artiodactyla, Prokaryotes to name just three.

The common theme in each case seems to me that we are lumbered with an historical taxon based on an 'obvious' extant division between two groups A and B, but then B turns out to be monophyletic within A, so suddenly A-excluding-B is paraphyletic, which makes it nonconforming to the desired taxonomical nomenclature that wants monophyly throughout.

Yes?
 
According to the latest edition of Dinosauria, Avialae, of which birds are a part, is a subtaxa of Eumaniraptora, which includes things like Deinonychus, Velociraptor, Troodon, and the like. This puts birds firmly within the reptile group. The exact position is likely to change once we get more fossils, however.

The common theme in each case seems to me that we are lumbered with an historical taxon based on an 'obvious' extant division between two groups A and B, but then B turns out to be monophyletic within A, so suddenly A-excluding-B is paraphyletic, which makes it nonconforming to the desired taxonomical nomenclature that wants monophyly throughout.

Yes?
There's two views on this. On the one hand, you have biologists, who generally want monophyly throughout all taxonomic nomenclature. On the other hand, you have paleontologists, who think that monophyly is great when you can get it, but frankly paraphyletic groups are incredibly useful in discussing the fossil record, particularly trends in evolution through time. The issue is that you need some way to separate the two groups as they diverge. If you're careful paraphytetic groups can give a lot of information.

There's a movement to switch taxonomy to a cladistic approach. Never got off the ground, though, thanks to the fact that it's very complicated. Systematics has the advantage of being easy to talk about.
 
"Systematics"

Is that what we call the non-cladistics version of taxonomy. And does it mean looking at the morphology and trying to pick out features to use as markers?
 
Thanks for that. Is their a simple term to describe the taxonomy that has Aves as a Class to distinguish it from the cladistic one?
Just "pre-cladistic classification" would get the idea across. I almost answered "Linnaean", because that's how the guy who invented the system categorized them, but someone would probably have argued that even with modifications such as extra levels being inserted and some species & groups being moved around, the system is still fundamentally Linnaean as long as it uses words like "class" and "order" and "phylum" and puts them in the right order.

I think I see the point you are making except that under the old-style taxonomy we were still able to call dinosaurs Reptilia, so it can't be completely correct that no account was taken of extinct species.
Extinct species can be fit into it after they're discovered, but that doesn't mean that it was set up with extinct species in mind. If it had been, then birds could never have been a class on their own. They only ended up as a distinct class in the first place because of the lack of anything else really similar to them to put in a group with them, and that lack of close relatives was due to extinctions.

It's the equivalent of all mammals but the cloven-hoofed ones going extinct. Someone (not a human of course) classifying living things in such a world (in the same kind of system as Linnaeus's) would have put the cloven-hoofed mammals as a class because they'd be different enough from all other living vertebrates (birds, reptiles, amphibians, & fish) to warrant being separated from them, and there'd be nobody else around to put in a group together with them. It would take the discovery of fossils of other things we now call mammals to change the cloven-hoofed animals' context and move their taxonomic level.

Really, the problem is that old Reptilia was paraphyletic because the historical taxonomists saw birds as being very different from reptiles, but internally consistent. Actually the latter is true, birds are monophyletic, but the former is not, if you exclude them from reptiles then old-style reptiles turn out not to be monophyletic.

Right so far?
Yes. But there's one more catch. To be sure that something is monophyletic, you have to not only check for fracturing within the group, as we've been dealing with so far in this case, but also check the larger group that it all fits within, see if there's any fracturing above as well as below. The "reptile" case happens to be a handy example of that.

Birds, non-avian dinosaurs, crocodilians, lizards, snakes, tuataras, pterosaurs, and ichthyosaurs all fit into a group called "diapsids". Mammals and some extinct critters (of which Dimetrodon is the most famous) make a group called "synapsids". Those two groups can be identified by the number of pits on each side of the head. (Yours are behind and a bit above the eyes, where the muscles you use in chewing are.) But there's one more group of amniotes (vertebrates that aren't fish or amphibians; they have amniotic eggs that don't need to be laid in water) I didn't mention: turtles. Their skulls lack pits, making them anapsids. And they're reptiles. So that would mean reptiles, even if you include all of the diapsids, still aren't necessarily monophyletic. Since it includes both diapsids and anapsids, a clade including them all must include the common ancestor of both, and all of that ancestor's descendants, which could very well mean including the synapsids (which means you've managed to narrow it down to all amniotes). Nobody knew at first which one of the three surviving amniote groups had split off from the other two first, before the remaining two split off from each other, so there were three possible arrangements, and under two of them, one of the two non-mammal groups could have ended up more closely related to mammals than to the other non-mammal group. For a while there, we seemed to have a two-thirds chance of getting one clade including some reptiles, and another one including mammals and the remaining reptiles.

As it turns out, that's all a could-have-been scenario. Biochemists found that turtle proteins & DNA look a lot like diapsid proteins & DNA, and could even match them to a particular group of diapsids (birds & crocodilians) more closely than to the others. This caused anatomists, whose discipline had said turtles were a distinct third 'apsid group in the first place, to say "Wait, did we miss something?" and recheck the anatomical work and come back with the answer "Yes, we did". It turned out that the pitless skull had been given too much "weight" in turtles' classification, and the weight of other clues in other parts of the body was consistent with a diapsid origin for turtles. Their skulls just smoothed out sometime after they split off from other diapsids. This mistake might have been influenced by the existence of fossils of true anapsids, a true third amniote group, which, since it doesn't include turtles after all, is entirely extinct.

So, even including all diapsids together, you still wouldn't necessarily have had a clade because of the anapsid issue, but, with turtles as archosaurs, it is. Also, even if turtles were anapsids, the question of whether they and diapsids together were a clade separate from synapsids would have been simply unanswered, until some more fossils or biochemical comparisons showed us the order of the splitting among those three clades. Then it would become known whether or not the group we're talking about had been a clade all along.

If Reptilia is the class containing birds, do we have a word that describes the paraphyletic group that would represent was we call reptiles in common language? Is there any use for such a word?
No. When there's already a common word that covers the meaning, and the established pattern is for scientific terms to apply to clades, there's no need for new scientific terms for groups that aren't clades.

"Systematics"

Is that what we call the non-cladistics version of taxonomy. And does it mean looking at the morphology and trying to pick out features to use as markers?
Yes, but in some cases human convenience & tradition set which morphological features are considered and which are ignored. For example, morphologically, we know that vultures come in two groups, each of which has some traits in common with some other group of non-vultures but not with the other group of vultures. But sometimes when we're talking about vultures, we really mean both groups of vultures and no non-vultures, so we're only concerned with traits that vultures have in common with each other and not with non-vultures. In that case, which morphological traits we decide matter and which ones we decide don't matter depends on the nature of the group we want to end up with.
 

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