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morphological characters in phylogenetic analysis...

Wulf Dec 02, 2004 03:10 AM

Hi folks,

I hope to learn something, so that I can understand

phylogenetic analysis usually are based on molecular data, such as mtDNA (e.g. cyt b gene). On the other hand, Wiens (2000a) describes the use of morphological data in phylogentic analysis mentioning different "ideologies" of character selection and character exclusion from the analysis.
Some taxonomists seem to exclude even any character that shows (even slightly) infraspecific variation, others code them as "?" (=unknown or variable) and even other taxonomists use them all. Wiens seems to support the use of characters showing infraspecific variation, too, but how reliable are the trees when using morphological characters showing infraspecific variation. I guess the phylogeny becomes even more tendable, right?

How would you (professional) guys i) select and ii) code morphological characters? What about the use of quantitative and qualitative characters? When it comes to body scales, are there rules for subdividing the characters (e.g. in Kluge 1993)

Character: Ventrals
240-252 ; coding 0
258-265 ; coding 1
270-275 ; coding 2 and so on...

I recognized a gap between the numbers of scales for the states.

Cheers,
Wulf

----
References:

Wiens, J.J. 2000a, In Phylogenetic analysis of morphological data, Smithonian Institution Press, Washington D.C.
-----
http://www.leiopython.de - the white-lipped python site -
http://www.herpers-digest.com - herp related eBooks search -

Replies (6)

CKing Dec 02, 2004 05:47 AM

Phylogenetic analysis is not "usually" based on molecular data. In fact, practically all such analyses were based on morphological characters prior to the 1960's. Even in the 1980's to early 1990's many analyses were still based on morphology alone. However, the ease with which DNA can be sequenced has made molecular systematics much more popular in recent years. The popularity of molecular systematics is understandable as well from the standpoint of reliability. Nevertheless, as I pointed out before, molecular data is not always available, especially in the case of fossils. Paleontologists therefore have to rely almost exclusively on morphological characters. That does not mean that such analyses are necessarily unreliable.

Dr. Michael Lee of the University of Queensland in Australia, for example, finds that snakes are closely related to varanoid lizards, and his morphological analysis has now been corroborated by Rest et al.'s (2003) mtDNA data. Dr. Lee is probably also correct that turtles are not diapsids in disguise because Rest et al.'s (2003) analysis showing turtles as diapsids is unreliable because of their choice of mammals as outgroup to the other amniotes. Mammals are amniotes too. Rest et al. assume (probably incorrectly) that mammals (actually synapsids) are the first group that branched off from the ancestor of the amniotes. Ironically, their tree then shows the placental and marsupials as two distinct lineages that form an unresolved polytomy with the lineage that is ancestral to all other remaining amniotes. That is of course anomalous because the placental-marsupial split occurred much more recently than the synapsid-anapsid split. Rest et al.'s tree shows the placentals and marsupials last shared a common ancestor at the same time as the synapsid-anapsid split!

What distinguishes Dr. Lee from most (but not all) other cladists is his willingness to analyze his characters. By doing so, he is able to achieve more reliable results than most cladists, who, in the words of Kurt Schwenk, "...cannot judge the quality of a character..." and must therefore merely "...assume that [all] characters are independent and hope that...enough 'good' characters will outweigh [the bad characters]". "Hope" is not a good tool for phylogenetic analysis; character analysis (e.g. comparative anatomy) is far better.

Yes, a good systematist like Dr. Lee can still come up with reliable results using morphological characters only. However, it is next to impossible to construct a reliable phylogeny using morphological characters alone. That is because lineages evolve at different rates morphologically. A lineage (e.g. the whales) can diverge so much from their close relatives that virtually all signs of their ancestry have disappeared. In such cases, morphological data often fails not only to unite closely related groups in an analysis, but morphological data often fails to reveal the paraphyletic nature of a given group. No one, for example, knew that Artiodactyla was paraphyletic prior to the use of molecular data, which shows that whales are descended from an artiodactyl that is closely related to the hippo, the pig and cattle. In fact, hippos are more closely related to whales than they are to the camel, another member of the Artiodactyla. Despite the fact that Dr. Lee correctly identifies varanoid lizards as the closest relatives of snakes, his analysis fails to show that the anguimorphs are a paraphyletic group.

In sum, morphological characters can be a reliable source of phylogenetic information, especially if their goodness is carefully analyzed (for example by using detailed comparative anatomy). Where morphological analyses often fail is their inability to delimit holophyletic groups. Many and perhaps most of the groups that the cladistic morphologists think are holophyletic often turn out to be paraphyletic or even polyphyletic. Hence it is not a surprise that even morphologists are turning more and more to the use of molecular characters when they try to ascertain branching order. Morphology nevertheless retains an important role in systematics and taxonomy. Morphological characters serve as corroborative evidence for molecular analyses. In taxonomy, it is next to impossible to classify organisms without studying their morphological disparity. A branching diagram alone, especially if it is based entirely on molecular data, is practically useless for delimiting both species and higher taxa.

Reference: Schwenk, Kurt 1994. Systematics And Subjectivity: The Phylogeny And Classification Of Iguanian Lizards Revisited. Herpetol. Rev. 25(2):53-57

PS Character choice, as well as character coding is a subjective process; different systematists see different numbers of characters when looking at the same part of the animal and they of course more likely than not code their characters differently.

Wulf Dec 02, 2004 12:37 PM

Hi CKing,

thanks for your comments. Indeed, I knew that phylogenetic analysis was based on morpholgical characters in earlier years, but since molecular analysis has become quite fashonable and perhaps more inexpensive it seems to be "state of the art" in noadays. The point I wanted to come to was if experiense has shown that phylogenetic analysis based on molecular data are more reliable than based on morphometric analysis. I do not want to say that the "old fashoned" workers were all wrong, but some papers that are based on molecular data support evidence for a different phylogeny of species than the ones created by using morphological data.
I understand that molecular data can not explain everything, as well as using only morphological characters. A combination of both would perhaps be the best choice. Well, often enough the results of molecular data and the resulting phylogenetic trees strongly differ from the ones produced by morphological data (and vice versa).

PS Character choice, as well as character coding is a subjective process; different systematists see different numbers of characters when looking at the same part of the animal and they of course more likely than not code their characters differently.

Subjective process, eh?
Well, that's interesting, and it's subject to discussions in the book edited by Wiens (2000a) mentioned in prior posting as well as to lots of papers from the same author. I have found some articles (online as PDF) that are quite interesting to read:

JOHN J. WIENS, 2001,Character Analysis in Morphological Phylogenetics:Problems and Solutions, Syst. Biol. 50(5):689–699
http://life.bio.sunysb.edu/ee/wienslab/wienspdfs/2001/characteranalysis2001.pdf

and others on Wiens web site:
http://life.bio.sunysb.edu/ee/wienslab/publicationpage.html

Cheers,
Wulf
-----
http://www.leiopython.de - the white-lipped python site -
http://www.herpers-digest.com - herp related eBooks search -

CKing Dec 02, 2004 09:43 PM

Wulf wrote:
"The point I wanted to come to was if experiense has shown that phylogenetic analysis based on molecular data are more reliable than based on morphometric analysis. I do not want to say that the "old fashoned" workers were all wrong, but some papers that are based on molecular data support evidence for a different phylogeny of species than the ones created by using morphological data."

Me:
Actually the great systematists of the past, who could not rely on molecular characters even if they wanted to, are not only not “all wrong,” but they are more likely to be correct than many modern day systematists who call themselves cladists. That is because these past greats often relied on detailed comparative anatomy, unlike many cladists who often rely uncritically on superficial similarities. The reason morphological characters are often unreliable is because they are often adaptive. If one reads Darwin’s book “Origin,” one can see how he and his colleagues often distrust adaptive characters and show more confidence in degenerate or “rudimentary organs.” If an organ is not needed by an organism’s way of life, Darwin reasons, then two organisms possessing the same rudimentary organ would be good evidence that they share a common ancestor. Darwin, in chapter 13 of “Origin”, also discusses at length the importance of embryology and ontogeny in phylogenetic analysis. Since embryological features such as gill slits are obviously not adaptive in mammalian embryos, their presence is thus strong evidence that mammals have an ancestor that possessed gill slits and hence gills. Mammals therefore once had an ancestor that breathed with gills. In contrast, most modern day cladists simply ignore phylogenetic clues that may be found in developmental biology, physiology or biogeography.

The reason molecular characters are often more reliable is because there are many parts of the genome of an organism that are more or less selectively neutral. The third nucleotide in each codon of a protein encoding DNA sequence, for example, can undergo mutation without affecting the amino acid being coded. Such mutations are therefore due entirely to chance. Two organisms with nearly identical nucleotide sequences, including the third nucleotides of all the codons, would almost certainly be closely related, since it would be unlikely that chance alone would produce identical random mutations in two different lineages evolving independently of each other. There are also molecules, such as serum albumin, which have no apparent function. These molecules are therefore selectively neutral and they are the analog of Darwin’s rudimentary organs. They too are often more informative of phylogenetic relationships than, say, ventral scale count, which is an adaptive character that can often evolve independently in two distantly related species.

Further, because morphology is more often adaptive, a species which has entered a new adaptive zone may undergo so much evolutionary change that its morphology may not resemble its closest relative. A classic example is that of the whales. They have no limbs and no hair and they are more fish like in their morphology than mammal like. Hence it was, until very recently, a mystery as to which group of mammal is ancestral to the whales. Despite the morphological disparity, the molecules of whales clearly show an affinity to the artiodactyls (camels, pigs, deers, cattle, hippos and relatives). Since most of the whales' molecules are not involved in the adaptation to the whales’ aquatic habitat, they retain their similarities to the whales' closest relatives, whereas almost every part of the morphology of whales have changed due to adaptation and nearly all morphological clues of the whales' ancestry have disappeared.

Morphological analyses therefore often fail to include divergent outgroups; such analyses result in the recognition of paraphyletic groups that the systematist may think, albeit falsely, are holophyletic. Phylogenies that are constructed on the basis of superficial, convergent similarities would result in unnatural polyphyletic groups. The morphological systematists who analyze their characters carefully often find themselves recognizing what turn out to be paraphyletic groups. The morphological systematists who think that they are "objective" by refusing to analyze the goodness of their characters would most likely find themselves recognizing polyphyletic groups.

Next time you read a morphological phylogenetic analysis, see if you can find any discussions at all of character goodness. If you cannot find any, then chances are that the results are not very reliable. Garbage in, garbage out applies equally to molecular systematics and morphological systematics.

johnscanlon Dec 03, 2004 01:18 AM

Wulf wrote....
>>
>>How would you (professional) guys i) select and ii) code morphological characters? What about the use of quantitative and qualitative characters? When it comes to body scales, are there rules for subdividing the characters (e.g. in Kluge 1993)
>>
>>Character: Ventrals
>>240-252 ; coding 0
>>258-265 ; coding 1
>>270-275 ; coding 2 and so on...
>>
>>I recognized a gap between the numbers of scales for the states.
>>
>>Cheers,
>>Wulf
>>
>>----
>>References:
>>
>>Wiens, J.J. 2000a, In Phylogenetic analysis of morphological data, Smithonian Institution Press, Washington D.C.
>>-----

If you're reading Wiens, that's the state of the art right there. There are lots of ways of coding morphological characters and he's tried most of them.

For a couple of other examples of morphological and combined analyses (shameless self-promotion here), try

Lee, M.S.Y. and J.D. Scanlon. 2002. Snake phylogeny based on osteology, soft anatomy and ecology. Biological Reviews 77(3): 333-402.

Scanlon, J.D., and M.S.Y. Lee. 2004. Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence. Zoologica Scripta 33: 335-366.

The latter paper (not yet posted, but should be up soon on David Williams' site, http://www.kingsnake.com/aho/pdf/pdfmain.html) may be of interest to some (Richard?) for discussion of historical biogeography, including Underwood's idea of Australian elapids descending from 'laticaudines'. Email me for a copy if it's not on the site yet.

Cheers,
-----
John D. Scanlon
Riversleigh Fossil Centre
Outback at Isa
Mount Isa, Queensland, Australia
riversleigh@outbackatisa.com.au

CKing Dec 03, 2004 09:53 AM

"If you're reading Wiens, that's the state of the art right there. There are lots of ways of coding morphological characters and he's tried most of them."

Me:
That is great except that he calls it "character analysis" instead of character coding. Character coding is a means of encoding a character so that it can be fed into the computer for phylogenetic analysis. The encoding process does not guarantee that a character is homologous, i.e. that it is a good character. The excellence of a systematic analysis depends on the goodness of the characters, not on the ability to encode taxonomic noise for analysis. I wish Wiens would spend more time developing new methods for eliminating phylogenetic noise instead of inventing new ways to encode it. Anyway, these methods mean little if the results are unreliable. Given the fact that morphological characters are often adaptive and given the fact that the rates of morphological change differ greatly within and among different lineages, morphological analysis will remain an inferior method as far as delimiting holophyletic groups (the holy grail of cladism) is concerned. Wiens' methods may be the last stand for the morphological cladists who are still holding out instead of switching to molecular systematics.

ScottThomson Dec 08, 2004 12:35 AM

CKing

"What distinguishes Dr. Lee from most (but not all) other cladists is his willingness to analyze his characters. By doing so, he is able to achieve more reliable results than most cladists, who, in the words of Kurt Schwenk, "...cannot judge the quality of a character..." and must therefore merely "...assume that [all] characters are independent and hope that...enough 'good' characters will outweigh [the bad characters]". "Hope" is not a good tool for phylogenetic analysis; character analysis (e.g. comparative anatomy) is far better."

ME

Yes I agree with this. I see that there has been two basic ideologies on the analysis and coding of morphological characters. The first is find everything you can asign it a number and crunch it. Gaffney used this in his paper on Turtle phylogeny of 1977 which placed Chelodina, Hydromedusa and Chelus as a monophyletic grouping.

Examination of the actual characters in detail, looking at development, function and their relationship to the bauplan of the turtles showed that none of these genera are closely related and that Hydromedusa was pretty much basal to the Chelidae, Chelus is closer to Phrynops and Chelodina is basal to the Australian radiation of the Chelidae. This was presented by Pritchard and received enormous condemnation for its "lack of objectivity".

Molecular analysis confirmed Pritchards view and demonstrated that Gaffney's phyllogeny was incorrect and was being effected by parallelisms. He did not predict that Long-necked piscivory could evolve on 3 occasions, in fact it may have evolved on four occasions. All the characters that united the "long-necks" in Gaffney 1977 were related to piscivory and required morphological adaptations for strike and gape mode hunting.

From my own research I no longer consider Chelus a long neck, though it has an appearance of being one. If one looks closely at the ratios of vertebrae length and shell length it actually has the same neck length as an equivalently sized Phrynops, but has a shorter shell. In other words its long neck is a bit of an illusion.

The reason I feel that molecular systematics can get a better tree than morphology when using methods similar to Gaffney is similar to what CKing was alluding to. The molecular sequence will be overpowered by the neutral genes due to the sheer number of characters used, whereas the morphology if not subjected to careful comparative anatomy will be overpowered by adaptive morphology. A good morphological character set with all characters justified by comparative anatomy can be as powerful a tool as a molecular one.

Cheers, Scott
-----
Scott Thomson

http://www.carettochelys.com
http://ittn.net

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