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dinoguy2
Joined: 22 Aug 2006 Posts: 41
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Posted: Tue 22 Aug, 2006 10:48 pm Post subject: Apomorphy-based names, crown groups, etc. |
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So, I was reading this paper (http://palaeo.gly.bris.ac.uk/donoghue/PDFs/2005/Donoghue_2005b.pdf) about crowns, stems, etc., and I got into some dangerous philosophical thinking that was messing with my mind a little, and I figured it'd help to write out my thought process.
A crown group, as we all know, is the clade containing the common ancestor of two living species. What got me thinking was the mention of things like tunicates, and how some researchers were terming them "stem chordates" and such. This got me thinknig about how crown groups are formulated--really, at least among amniotes, all the crown groups correspond almost exactly to the contents of the traditional linnean classes.
For instance, crown group Aves. Why were Passer + Struthio chosen as specifiers rather than, say, Passer + Crocodilus? Obviuosly, because crocodiles are tradionally "reptiles" and ostriches are traditionally "birds". Essentially, defining any group based on extant species is just as arbitrary as Linnean taxonomy becasue it's doing the exat same thing, with a veneer of phylogenetic nomenclature slopped over it.
The telling implication here also is that vernacular terms are directly linked to clade names, especially for crown groups. Why does "bird" = Aves? (Because "aves" *means* "bird"!) Better yet, why was the loaded name Aves even *used* in phylogenetic taxonomy?
If, for instnace, an animal is found to fall within the clade Chordata, that lacks a notochord... well, it wouldn't be the first time the name of a group istelf was misleading as to that group's contents, but still.
I think this whole PN, cladistics, vs oldschool Linnean situation we have now is that much more messy because of the actual nomenclature involved. PN should never have adopted the names of former Linnean groups. Names should be descriptive, especially in PN and cladistics. What's the utility of naming a clade Aves, which implies a very specific suite of apomorphies right in then ame, instead of something like "Passerarchaeopteryx"? Or simply "CrownPasserStruthio" or "StemPasserArchaeopteryx"?
Why not limit the traditional linnean names to something that more accurately refelcts their nomenclature, such as apomorphies? Already we name node- and stem-based clades that may or may not be redundant with each other. Why not have Aves = "Primary feather w/ a central vane as in Passer", Chordate = "Notochord as in Homo", etc. Cladistic methodology, and PN in general, seems always bogged down because it's trying to teach an old Linnean dog new tricks. Why not leave the old names, or at least modify them with definitions matching the contents they describe, and use new names for a new system?
Would certainly fix problems regarding "standard" endings like -idae, at least. |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Wed 23 Aug, 2006 2:30 am Post subject: Re: Apomorphy-based names, crown groups, etc. |
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| dinoguy2 wrote: | | A crown group, as we all know, is the clade containing the common ancestor of two living species |
Well, it's the clade consisting of the final common ancestor of two or more extant specifiers (species or specimens), plus all descendants of that ancestor.
| dinoguy2 wrote: | | What got me thinking was the mention of things like tunicates, and how some researchers were terming them "stem chordates" |
Eek.
| dinoguy2 wrote: | | really, at least among amniotes, all the crown groups correspond almost exactly to the contents of the traditional linnean classes. |
I disagree. Only two of the three traditional amniote classes (Aves and Mammalia) correspond (roughly) to crown groups. Classis Reptilia as generally used is doubly paraphyletic. And none of the other traditional vertebrate classes really correspond to crown groups. (You could make cases for Chondrichthyes and Amphibia, although those are far from clear-cut. And Osteicthyes and Agnatha are right out.)
| dinoguy2 wrote: | | If, for instnace, an animal is found to fall within the clade Chordata, that lacks a notochord... well, it wouldn't be the first time the name of a group istelf was misleading as to that group's contents, but still. |
Nobody has a problem with snakes being tetrapods; why not a secondarily notochord-less chordate?
A more likely case would be the discovery of an animals with a notochord lying outside of crown chordates. But they could be referred to as "apo-chordates" (or whatever the final form of that notation is). Similarly, there were undoubtably non-mammalian theropsids with lactation; these would not be mammals, but could be termed "apo-mammals".
| dinoguy2 wrote: | | PN should never have adopted the names of former Linnean groups. |
I've thought the same thing myself sometimes, but it isn't realistic. You'd be solving one problem but creating a bigger one: namely, that people would have to learn an entirely new set of names.
| dinoguy2 wrote: | | Why not limit the traditional linnean names to something that more accurately refelcts their nomenclature, such as apomorphies? |
It isn't always clear which apomorphy a name refers to, if any.
| dinoguy2 wrote: | | Why not have Aves = "Primary feather w/ a central vane as in Passer" |
Case in point.
| dinoguy2 wrote: | | Chordate = "Notochord as in Homo" |
I actually don't have a huge problem with this one, although it seems like there should be some way to refer to the crown clade. And if well-known names are used everywhere else for crown clades, then this would actually be (strangely enough) counterintuitive. _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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dinoguy2
Joined: 22 Aug 2006 Posts: 41
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Posted: Wed 23 Aug, 2006 3:08 am Post subject: |
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| Quote: | | Nobody has a problem with snakes being tetrapods; why not a secondarily notochord-less chordate? |
Well I meant a *primarily* notochordless chordate, which I admit is highly unlikey (given current definitions i guess the notochord would have to have arisen twice independantly at the base of the group?).
| Quote: | | A more likely case would be the discovery of an animals with a notochord lying outside of crown chordates. But they could be referred to as "apo-chordates" (or whatever the final form of that notation is). |
Much better example. Why develop a new term like apo-chordate when a perfectly good term for "notochord as in insert-favorite-vertebrate-here" already exists? Why not invent a new term for the clade amphioxus+vertebrates, or however Chordata is defeined phylogenetically, rather than use the old term for the (approximate) old definition, and a new term for the new one if it describes a different group?
| Quote: | | You'd be solving one problem but creating a bigger one: namely, that people would have to learn an entirely new set of names. |
Well, not entirely new. If the apomorphy-defined 'Linnean' groups (sans ranks, naturally, when used in a phylogeny) are reduntant with the new ones, use those. Chordata would have priority over New-name-ata so long as the content of the apo-clade matches that of the node-based clade (as currently done when you have two node- and stem-based clades which, at the moment, describe the same content). Besides, the advent of cladistics means that names will proliferate anyway--not that that's a bad thing, and i know every clade in the world doesn't need a name, but what's learning a few extra/replacement names gonna hurt?
| Quote: | | It isn't always clear which apomorphy a name refers to, if any. |
In these cases, if no apomorphy could be found at all to unite even the traditional contents, chuck it out with Vermes and Reptilia. Or incorporate the old name into the replacement for a bit more nomenclatural stability if you're so inclined.
As for Aves, which refers to a whole suite of apomorphies, I just picked the most obvious. Nobody says these definitions have to match exactly traditional uses, but unless kiwi are a lot more primitive than we think, it's certainly not going to exclude the crown group.
| Quote: | | And if well-known names are used everywhere else for crown clades, then this would actually be (strangely enough) counterintuitive |
How so? |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Wed 23 Aug, 2006 4:03 am Post subject: |
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| Quote: | | Quote: | | And if well-known names are used everywhere else for crown clades, then this would actually be (strangely enough) counterintuitive |
How so? |
If the practice of converting well-known, long-used names to crown clades is widely followed, I would come to simply expect it. I would simply expect Insecta, Angiospermae, Archosauria, etc. to be crown clades, and would be surprised to discover that Chordata was treated differently.
Of course, you could use apomorphy-based definitions for these names (except for cases where the etymology does not derive from an apomorphy, e.g., Insecta, Aves, etc.), but to be consistent you'd have to do it across the board. And knowing how many pitfalls there are with apomorphy-based definitions, it seems ill-advised to give them to all the best-known names out there.
Consider if we did it to Mammalia. We have absolutely no idea when lactation began within Theropsida. It would possibly make cynognathids mammals, and might even include things like therocephalians and dicynodonts. It's even conceivable (if barely so) that such a move could make Dimetrodon a mammal! In any case, an apomorphy-based definition would make the name extremely difficult to apply. While I'm not necessarily against definitions that are hard to apply, it doesn't seem wise to give them to well-known names.
I think it's simpler to let Mammalia be a crown clade, and then discuss whether or not various fossil theropsids (or "pan-mammals") are "apo-mammals". _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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dinoguy2
Joined: 22 Aug 2006 Posts: 41
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Posted: Wed 23 Aug, 2006 5:06 pm Post subject: |
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[quote="tmkeesey"] | Quote: | In any case, an apomorphy-based definition would make the name extremely difficult to apply. While I'm not necessarily against definitions that are hard to apply, it doesn't seem wise to give them to well-known names.
I think it's simpler to let Mammalia be a crown clade, and then discuss whether or not various fossil theropsids (or "pan-mammals") are "apo-mammals". |
Well, if Mammalia is based on lactation, which it has always traditionally been (isn't that why Linneaus included bats in mammalia rather than aves?), I don't see a practical problem with having that high degree of uncertainty with fossil species. Talking about "Mammalia" would in effect be talking about lactating synapsids, the only ones we can be confident of being extant and a very few extinct forms. If we don't know that such and such extinct animal is a mammal or not, it probably is not going to change the way we talk about that group and its apomorphy.
Placing the tag "mammalia" at an arbitrary point that *definitely* leaves out lactating synapsids is in my opinion worse, because the name itself implies this meaning. The crown group should be called something else, like corona-Mammalia, so that your explicit in you inclusion only of lactating synapsids that are presently extant. This could, actually, be even more useful if it was defined as such--that is, have "extant" itself be an apomorphy!
Then, if monotremes go extinct, they automatically remove themselves from the group. Discussing a group meant to represent only extant forms would get pretty messy if presently exinct forms were included by definition...  |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Wed 23 Aug, 2006 6:05 pm Post subject: |
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Etymology does not and should not drive nomenclature. Dinosauria and Basilosaurus are not lizards, synapsids do not have fused fenestrae derived from the diapsid condition, and Pongo pygmaeus is actually quite large.
Of course we can make some concessions to etymology, but we shouldn't bend over backwards. Equating Mammalia with lactation would be bending over backwards. It would make the clade extremely unstable, perhaps nearly as inclusive as Theropsida, perhaps nearly as exclusive as the crown group.
Unstable apomorphy-based definitions may have their place, but not, I think, for such common, well-known, often-used names as Mammalia.
I do agree that there could be a notational prefix for crown groups; perhaps you could say that mammals are corono-therapsids, or that Mammalia = crown(Therapsida). _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Wed 23 Aug, 2006 6:21 pm Post subject: |
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| dinoguy2 wrote: | | This could, actually, be even more useful if it was defined as such--that is, have "extant" itself be an apomorphy! |
I agree that "extant" is (more or less) a chronological apomorphy.
| dinoguy2 wrote: | Then, if monotremes go extinct, they automatically remove themselves from the group. Discussing a group meant to represent only extant forms would get pretty messy if presently exinct forms were included by definition...  |
For an apomorphy-based definition, the apomorphy is useless unless it is:- Fixed
- Absolute
...by which I mean it cannot change arbitrarily under various circumstances (e.g., different times), and every organism must either possess or lack it definitively.
Others have suggested defining "extant" in the PhyloCode as something along the lines of, "recent enough that museums possess specimens from living organisms", which I think nicely cuts to the quick of the matter and satisfies the conditions. (This may be a topic for a new discussion in the PhyloCode board....)
But, apart from that, I think it's generally agreed that, in most instances, crown clades should be defined using node-based definitions that happen to have extant specifiers. Thus, the word "extant" is not a necessary part of a crown clade definition. Mammalia can simply be defined as the node-based clade specified internally by Ornithorhynchus anatinus and Homo sapiens. Those species happen to be extant, so it's a crown clade.
Thus, crown clades can be used as anchors for the "pan-" and "apo-" formulas. If X is a crown clade defined using a node-based definition, then "pan-X" means "the branch-based clade specified internally by the internal specifiers of X and externally by all extant nonmembers of X". Similary, if the name "X" refers to apomorphy "M", then "apo-X" would mean, "the apomorphy-based clade specified by the internal specifiers of X and apomorphy 'M'." _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc.
Last edited by tmkeesey on Thu 24 Aug, 2006 5:31 am; edited 1 time in total |
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dinoguy2
Joined: 22 Aug 2006 Posts: 41
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Posted: Wed 23 Aug, 2006 10:06 pm Post subject: |
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TMk said in the same discussion on another board: | Quote: | Etymology does not drive nomenclature. You can have bipedal or even limbless tetrapods, dinosaurs that or neither "fearfully great" nor lizards, and archontans that totally do not rule. Pongo pygmaeus is actually gigantic, Basilosaurus is not even a reptile let alone a lizard, and Homo sapiens is frequently foolish.
Names are just names. |
And I think they should be more. Wouldn't it be more useful to have names that mean something and convey information, rather than names that mean diddly squat? With PhyloCode, we have a great opportunity ot improve the way things are done, not just change them around.
| Quote: | | Thus, crown clades can be used as anchors for the "pan-" and "apo-" formulas. if X is a crown clade defined using a node-based definition, then "pan-X" means "the branch-based clade specified internally by the internal specifiers of X and externally by all extant nonmembers of X". Similary, if the name "X" refers to apomorphy "M", then "apo-X" would mean, "the apomorphy-based clade specified by the internal specifiers of X and apomorphy 'M'." |
Actually, I do kind of like this setup. I assume "the apomorphy-based clade specified by the internal specifiers of X and apomorphy 'M'." is equivilant to "apomorphy M as in specifer X1 and X2"?
I'd still be a little miffed that we need to prefix the apo-clades for groups that were originally based on apomorphies in the first place, but it's usefulness is the same either way. |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Wed 23 Aug, 2006 11:29 pm Post subject: |
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| dinoguy2 wrote: | | Actually, I do kind of like this setup. I assume "the apomorphy-based clade specified by the internal specifiers of X and apomorphy 'M'." is equivilant to "apomorphy M as in specifer X1 and X2"? |
Well, I'd phrase it as, "the initial common ancestor of the internal specifiers of X to possess trait 'M' homologous with trait 'M' in the internal specifiers of X, plus all descendants of that ancestor." In this case, "the initial common ancestor of Ornithorhynchus anatinus and Homo sapiens to possess lactation homologous with lactation in Ornithorhynchus anatinus and Homo sapiens, plus all descendants of that ancestor."
| Quote: | | I'd still be a little miffed that we need to prefix the apo-clades for groups that were originally based on apomorphies in the first place, but it's usefulness is the same either way. |
Quite, and this way we get to have a (relatively) stable Mammalia. Never mind arguments about whether some eucynodonts belong or not; it's still far more stable than apo-Mammalia. And it can be applied under any phylogenetic hypothesis, while apo-Mammalia requires additional (and probably irretrievable) knowledge for exact application. _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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David Marjanović
Joined: 24 Aug 2006 Posts: 137 Location: currently Paris
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Posted: Thu 24 Aug, 2006 12:50 pm Post subject: |
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Hey, dinoguy2! Good to see you here! That's easier than discussing on a Wikipedia talk page
I think that apomorphy-based definitions should be used for rather few clades -- those with apomorphies for which we don't need to expect lots of transitional states, and those for which anything but an apomorphy-based definition could result in drastic changes in concept. The turtles (Testudinata sensu Joyce, Parham & Gauthier 2004) fulfill the first criterion: the shoulder blades can be either inside or outside the ribcage. (At least for amniotes, which always have long ribs.) Diapsida fulfills the second: if it gets a branch-based definition, we can discover anapsid diapsids ( = diapsids with an anapsid skull roof), and if it gets a node-based one, we can discover diapsid non-diapsids. Diapsida does not quite fulfill the first condition, though, so a careful choice and delimitation of the apomorphy is required here. I suggest the upper temporal fenestra -- in the right place (ventrally delimited by postorbital + squamosal), larger than a foramen, and with rounded, smooth edges unlike a fontanelle (Milleretta has a lower temporal fontanelle, for example). Tetrapoda may fulfill both conditions, especially if the digital arch hypothesis is correct. (It clearly fulfills the first, in any case!)
In general, however, I don't recommend apomorphy-based definitions. Tetrapoda, for which I still recommend an apomorphy-based definition because of all the above, will illustrate the principal problem: We don't know if Ventastega, Elginerpeton, and several other taxa had digits and/or fin rays, so we don't know if they are tetrapods, even though their phylogenetic position is fully resolved! Anchoring Mammalia on lactation, for example, would be downright foolish.
For Mammalia the presence of the secondary jaw joint (which is not correlated to the absence of the primary one!) could still be used, but this makes Haramiyidae, Tritylodontidae, and Tritheledontidae mammals (they were usually excluded). A node-based definition, such as the one anchored on Sinoconodon (Luo, Cifelli and Kielan-Jaworowska 2002) or the crown-group one, may be a better idea.
For Aves? Forget it.
For Amniota? Anything but the amniotic egg. Eggs fossilize very, very rarely (unless they have a hard shell -- but that's not the normal condition). We don't know how Diadectomorpha and Solenodonsaurus or the possibly relevant Westlothiana and Casineria reproduced, so we wouldn't be able to tell if they are amniotes! Everyone uses Amniota for the crown-group, so let it be that crown-group.
Addition after preview: This forum does not even take en dashes. There are outright Unicode fora out there... |
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dinoguy2
Joined: 22 Aug 2006 Posts: 41
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Posted: Thu 24 Aug, 2006 3:20 pm Post subject: |
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| David Marjanović wrote: |
In general, however, I don't recommend apomorphy-based definitions. Tetrapoda, for which I still recommend an apomorphy-based definition because of all the above, will illustrate the principal problem: We don't know if Ventastega, Elginerpeton, and several other taxa had digits and/or fin rays, so we don't know if they are tetrapods, even though their phylogenetic position is fully resolved! Anchoring Mammalia on lactation, for example, would be downright foolish. |
This is actually why I DO advocate using apo-clades, but parallel to traditional phylogenetic definitions. There are some things about the fossil record we are simply never going to know. We probably will *never* know where lactation began in synapsids. That doesn't make this group useless to discuss, it just means the basal membership will always be fuzzy. So what? If you want to talk about a group of apomorphies, it's implicit that you mean a group composed of organisms *known* to have this. If you want to talk about a group based on descent, that's what PN is for (not that apomorphy-based groups are not within the realm of PN, but you know what I mean).
In the spirit of the argument i'm making in this thread, i'd oppose naming an apo-Mammalia based on the characteristic jaw joint, as this is not conveyed in the name of the clade. Mammalia refers specifically to the mammary gland.
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For Aves? Forget it.
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Why? I know it's based on an apomorphy suite, but just come to a consensus on one and go with it. Or better yet, use the suite of apomorphies in the definition. Don't tell me science isn't about consesnus
| Quote: | | wouldn't be able to tell if they are amniotes! Everyone uses Amniota for the crown-group, so let it be that crown-group. |
Well, then use your best guess based on their phylogenetic position, and list them as possible amniotes, probable amniotes, possible non-amniotes, or whatever. What's wrong with admitting "we knoe they're within clade X but we can't be sure whether or not they were true amniotes."? |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Thu 24 Aug, 2006 5:45 pm Post subject: |
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| David Marjanović wrote: | | Addition after preview: This forum does not even take en dashes. :shock: There are outright Unicode fora out there... |
The terminal "ć" in your username is also improperly rendered in some places.
Well, that's open source for you. I'll look into it when I have some more time.... _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Thu 24 Aug, 2006 5:53 pm Post subject: |
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| dinoguy2 wrote: | | This is actually why I DO advocate using apo-clades, but parallel to traditional phylogenetic definitions. |
By "apo-clade" do you mean the "apo-" prefix convention, or "apomorphy-based clade"? ( I assume the latter, but I think the term should be used for the former.)
| dinoguy2 wrote: | | In the spirit of the argument i'm making in this thread, i'd oppose naming an apo-Mammalia based on the characteristic jaw joint, as this is not conveyed in the name of the clade. Mammalia refers specifically to the mammary gland. |
Definitely.
"Apo-Mammalia" would not have to be specifically named, though--"apo-" could just be an informal (or semi-formal or whatever) prefix, like "pan-" or even "non-".
| dinoguy2 wrote: | | David Marjanović wrote: | | For Aves? Forget it. |
Why? I know it's based on an apomorphy suite, but just come to a consensus on one and go with it. Or better yet, use the suite of apomorphies in the definition. Don't tell me science isn't about consesnus  |
Using the full suite will just give you the crown clade (or indistinguishably close to it), anyway, so why not use a simple node-based definition instead of listing dozens of apomorphies (none of which are apparent from the name itself)?
| dinoguy2 wrote: | | Well, then use your best guess based on their phylogenetic position, and list them as possible amniotes, probable amniotes, possible non-amniotes, or whatever. What's wrong with admitting "we knoe they're within clade X but we can't be sure whether or not they were true amniotes."? |
Or, what's wrong with saying, "Diadectes is not an amniote, but it may be an apo-amniote (and is definitely a pan-amniote)"? _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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David Marjanović
Joined: 24 Aug 2006 Posts: 137 Location: currently Paris
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Posted: Thu 24 Aug, 2006 11:31 pm Post subject: |
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Is there more than one font in this forum? That would explain it very easily.
I wouldn't say that Aves refers to a suite of apomorphies. It's just the Latin word for "birds", and the definition of that is "I know it when I see it". You can try to come up with a phylogenetic definition that keeps the content of this largely intact; I have proposed such a definition on both meetings and hope to publish it sometime. But I can't think of even just one apomorphy that would reliably diagnose such a clade. |
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tmkeesey Site Admin
Joined: 14 Aug 2006 Posts: 287 Location: Los Angeles, CA, USA
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Posted: Fri 25 Aug, 2006 1:58 am Post subject: |
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| David Marjanović wrote: | | Is there more than one font in this forum? That would explain it very easily. |
That would make the character show up as a block or a question mark, not as an HTML entity. It's almost certainly more complicated than that.
We're just using the phpBB default settings, so....
| David Marjanović wrote: | | I wouldn't say that Aves refers to a suite of apomorphies. It's just the Latin word for "birds", and the definition of that is "I know it when I see it". |
I quite agree. "Apo-avian" would be a completely useless term (one peculiarity of the "apo-" prefix). _________________ T. Michael Keesey
Director of Technology, Exopolis, Inc. |
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