Such expressions as that famous one of Linnæus, and which we often meet with in a more or less concealed form, that the characters do not make the genus, but that the genus gives the characters, seem to imply that something more is included in our classification, than mere resemblance. I believe that something more is included; and that propinquity of descent,—the only known cause of the similarity of organic beings,—is the bond, hidden as it is by various degrees of modification, which is partially revealed to us by our classifications (Darwin, 1859, p. 413f).
Showing posts with label Haeckel. Show all posts
Showing posts with label Haeckel. Show all posts

Thursday, 29 November 2007

Natural and Artificial Classification: A reply to Wilkins

The following post is a reply to John Wilkin’s The philosophy of classification on his blog Evolving Thoughts.

An Uninformed Consensus

John Wilkins in his recent post believe that our view is "radical" because
"… they have presented some views on classification that do, indeed, differ from the received consensus."
We beg to differ.

In late 20th and early 21st century literature there are very few discussions on the nature of classification. Most revolves around explaining existing classifications (i.e. Reptilia) or in the defence of poorly defined taxonomic groups that fail to form groups (i.e., paraphly). It is these debates (i.e., paraphly versus monophyly) that would benefit from the discussions of early 20th and late 19th century morphologists, would did hold a consensus view of natural and artificial classifications. That consensus was this,
We then follow a Natural Method, which cannot be called a system, because it is destitute of any unity of principle. (Candolle & Sprengel, 1821)
It is our belief that the pursuit for explanations to existing classifications that ended this debate and therefore any consensus. Furthermore, it is the addition of homology = similarity that radically altered how we view classifications, leading to the almost Fukuyamaist statement that,
"I would say that the effort put into this controversy is further evidence that systematists do not have their priorities straight. In their day-to-day work they really do not make much use of classifications, but they show a strange obsession with fighting about them for reasons that seem to me to be an historical curiosity" (Felsenstein 2005)
Currently there is no consensus over natural or artificial classifications. The topic is a moot point and very few concern themselves with its relevance to 21st systematics and biogeography. As systematists we are more or less tied to the consensus of the past, namely to the literature of the 19th century and early 20th century. In that sense we are not “radicals", but rather “old fashioned”.

Similarity and Homology

Similarity, as expressed in the usual kinds of data matrices, is 11, or, the molecular version, AA is not a relation. The 11 and the AA are, if anything, homologues, the parts, the 'namesakes' as Owen called them. We see homology as a relation: 0(11), or the molecular version, G(AA). We stated earlier:
"...all molecular systematic studies are phenetic as they ignore relationship, that is, homology". One might expand that and say, "...all numerical systematic studies are phenetic as they ignore relationship, that is, homology."
This would be more accurate.

In response to John’s comment,
"I'm not sure I follow this. According to current usage, molecular systematics does rely on homologies: they have a number of special terms devoted to identifying them: paralogy, xenology and orthology. Of course, they often don't use homology properly. And to identify a homology in molecular biology you need to do some prior work; homology is an inference from sequence similarity (including eyeball alignment). In short, if I understand the argument, molecular systematics derives homology from similarity".
In fact we would suggest that it would be more accurate to say:
"... molecular systematics does rely on HOMOLOGUES: they have a number of RELATIONS DERIVED FROM them: paralogy, xenology and orthology....And to identify a HOMOLOGUE in molecular biology you need to do some prior work; HOMOLOGUES ARE inferenceS from sequence similarity (including eyeball alignment). In short, if I understand the argument, molecular systematics derives HOMOLOGUES from similarity ..."
This certainly is not radical. What we are suggesting is that de Candolle (1813) presented a very clear account of classification, an account still of significance today.

Haeckel and Classification

In our understanding, Ernst Haeckel did more than most to promote the genealogical view of species relationships. It might be fair to say that all our genealogical endeavours stem from Haeckel. Adolf Naef (1917, 1919)was the first to critique that viewpoint His interest was in natural classification. Hennig (1950), quite deliberately, focused on Naef. Thus, it might be fair to say that Hennig's efforts were directed towards rehabilitating Haeckel. Further, one might see Systematics and Biogeography (Nelson & Platnick, 1981) as a further detailed critique of Haeckel - if the most detailed critique available - and a restatement of de Candolle's viewpoints on classification. In this sense cladistics sensu Nelson & Platnick is of greater significance than cladistics sensu computer programs.

We would venture the suggestion that Sober (1988) mistook cladistics sensu Farris (parsimony sensu Farris) as if it was the generally accepted view (in the mid-1980s that might have been possible). In fact Sober deliberately excludes the more general view, as if the argument really was about parsimony versus likelihood, one algorithm versus another,
"Because this work is about phylogenetic inference, not classification, nothing will be said about the current controversy concerning so-called 'pattern' cladism." (Sober, 1988:8, footnote 7).
Thus, in our view, the more general study of classification exclude Sober's work as a relevant commentary on the matter.

References
Candolle, A.P., de, & Sprengel, K. 1978. Elements of the philosophy of plants. Reprint of the 1821 ed.. New York, NY.
Hennig, W. 1950. Grundzüge einer Theorie der phylogenetischen Systematik, Deutsche Zentralverlag, Berlin.
Naef, A. 1917. Die individuelle Entwicklung organischer Formen als Urkunde ihrer Stammesgeschichte: (Kritische Betrachtungen über das sogenannte "biogenetische Grundgesetz"), Verlag von Gustav Fischer, Jena.
Naef, A. 1919. Idealistische Morphologie und Phylogenetik (zur Methodik der systematischen), Verlag von Gustav Fischer, Jena).
Nelson, G. & Platnick, N.I. 1981. Systematics and biogeography. Cladistics and vicariance. Columbia University Press, New York.
Sober, E. 1988. Reconstructing the Past: Parsimony, Evolution, and Inference. MIT Press, Cambridge, Massachusetts.

Monday, 5 November 2007

Haeckel, Hennig and History: Evolving Thoughts and Words

John Wilkins, in his eminently readable and ever provocative blog Evolving Thoughts, presents an account of some historical matters relevant to Natural and Artificial Classifications, matters that might illuminate the differences of opinion between Joe Felsenstein and ourselves. To be sure, we differ on certain fundamental matters. But the issue of natural classifications is a subject that might repay closer attention and discussion. John's history is a cast of worthy individuals (Adanson, Linnaeus, Agassiz, Macleay, etc.), many who made worthy contributions to discovering the means with which to discover natural classification. They all, to one degree or another, had some sort of interpretation of that classification. They all, to one degree or another, had some sort of axe to grind. Never mind.

John moves on to note that "...it is with Haeckel and the early German paleontologists of the 20th century that phylogenetic relations become the core of classification, and we all know, of course, that Hennig defined a natural group as a monophyletic group." Haeckel is a something of a departure and one we see of significance. Here's Agassiz on Haeckel:
"It is not that I hold Darwin himself responsible for these troublesome consequences. In the different works of his pen, he never made allusion to the importance that his ideas could have for the point of view of classification. It is his henchmen who took hold of his theories in order to transform zoological taxonomy" (Agassiz, 1869: 375, our translation) (see also http://www.athro.com/general/atrans.html.
Those henchmen included Haeckel. Most of Haeckel's genealogical trees ('phylogenies') were linear schemes of hypothesized relationships, with some taxa 'giving rise' to others, that is, paraphyletic groups not so much created by him (many were, of course) but retained and explained in terms of ancestry and descent, in terms of evolutionary relationships, relative to a particular model of change. It was to Hennig's credit that Haeckel's paraphyletic groups were exposed for what they were: empty conventions. And thus, a circle was closed and certain groups understood as not part of the discovery process of classification - or so it seemed. Haeckel's problem was taking a viewpoint (ancestry and descent) and interpreting classification from that perspective.

Now that's not a whole million miles away from the current viewpoint:
find a model of evolution and interpret the data from that point of view. Still, again, never mind. What comes shining through most of the earlier contributions to the debate is that, one way or another, Adanson, Linnaeus, Agassiz and Macleay, among others, did have a notion of the centrality of classification: homology. So when John suggests that "...It does not seem to me that cladistic classification is in possession of a notion of taxon that grounds its classifications" he omits consideration of homology. But he is not alone. The entire crop of books recently published dealing with the 'mathemetisation' of phylogeny do not deal with that subject at all. Thus, or so it seems to us, the 'mathemetisation' of classification (phylogeny) has lead to a profusion of artificial methods.

We finished a recent (as yet unpublished) paper with the following words, the first few are from Gareth Nelson:
'"What, then, of cladistics in relation to the history of systematics? If cladistics is merely a restatement of the principles of natural classification, why has cladistics been the subject of argument? I suspect that the argument is largely misplaced, and that the misplacement stems, as de Candolle suggests, from the confounding goals of artificial and natural systems" (Nelson, 1979, p. 20). Cladistics is concerned with homology, monophyly, evolutionary patterns, taxa (species), and natural classifications. That is, natural classification is concerned with relationships.'
PS. One of us [DMW] is a Londoner. We are aware (sometimes painfully) of the relationship between Australia and the UK capital city, the latter a onetime plentiful source of persons to inhabit the former. The 'relationship' between Australian's and Londoner's is such that when travelling in the United States a London accent is often mistaken for an Australian accent. We mention these facts, partly because the other half of this pair [MCE] is an Australian. And partly because the word 'Barny' is also said to originate from Cockney Rythming slang, as in Barny Rubble = trouble, and thus (if true) a mingling of Australian slang, London slang and American cartoon characters - words really do have a life of their own!