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26 March 2004
Friday arachnid blogging, and what's in a name
After last week's grand-guignol, here's a more pastoral scene altogether:
The orangey spider tucked in among the purple petals is one of the 'running spiders' or, more properly, philodromid crab spiders. She is very likely Philodromus rufus though, as she is a juvenile, it is extremely difficult to identify her precisely.
The name 'running spider' might seem odd. After all, most spiders (at least those that do not spend their entire lives on a web) run. But in this case the popular name derives from the Linnaean name Philodromus, 'one who loves to run', and they are quick little things indeed. What is really interesting here is the term 'philodromid crab spider'. They are so called to distinguish them from the thomisid crab spiders. Remember that terminal -d, by the way, because it's going to be important.
If you've gone to look at the thomisids on Ed Nieuwenhuys's excellent website and compared them to my young philodromid, you'll probably have noted that they look very much alike, save for one detail. The thomisids have stubby little hind legs (pairs 3 and 4) while their two forward pairs of legs, especially the first pair, reach heroically Popeye-like proportions. The forward legs of the philodromids are very similar, but their hind legs aren't nearly as stubby; in fact they look more or less 'normal'.
Now, before we contemplate drawing any conclusions about the degree of relation between thomisids and philodromids, we can explain the difference in leg pattern by the respective lifestyles of the two sorts of spider. Philodromids chase down their prey. Thomisids sit stock-still on a plant, often in the middle of a flower, and wait until an insect walks into the grasp of their prodigious forearms; then wham and it's lunchtime.
People used to say 'Well, we have crab spiders; clearly this one sort are different to the other sort, so we'll give them different names, but they're essentially part of the same group'. And so we had all of these spiders in the family Thomisidae, with the short-rear-legged lot called Thomisinae and the longer-rear-legged lot called Philodrominae. The -idae ending denotes a family (one level up from genus in the classical hierarchy), while -inae denotes a subfamily.
What you need to know about taxonomists (people who classify organisms) is that they fall into one two big groups, the lumpers and the splitters. As the names suggest, lumpers are inclined to shove disparate organisms into a single group if they have any shared characteristics; whilst splitters are, shall we say, much given to celebrating diversity.
As far as the crab spiders go, the splitters have tentatively won the day. Today one no longer speaks of Philodromines but of Philodromids. The d makes all the difference; the spider in the picture above is not part of a sub-goup of Thomisidae but rather member of an independent family in its own right, the Philodromidae.
This might strike you as so much hair-splitting, but it isn't. Almost all taxonomists these days have adopted a cladistic worldview. Cladistic systematics is notorious for rigidly dogmatic insistence on an essentially Martian terminology, but brush aside the bizarre words and it's pretty intuitive and (I think) sound: if you want to divide up the world's organisms, the terms you use should reflect the actual evolutionary history of those organisms.
To use a concrete example: as we've all heard, 'birds are really dinosaurs'. This might strike you as odd, given that (i) dinosaurs, as every shoolchild knows, became extinct a squillion years ago and (ii) that thing on your plate is a chicken, for chrissake, not a dinosaur. But to a cladist (and we are all cladists now), the point is that no taxonomic group is 'real' unless it contains all the organisms descended from a single common ancestor, and no organisms not descended from that ancestor. Birds as we know them today arose within one particular group of reptiles; that group was the dinosaurs. So a Tyrannosaurus and a chicken are more closely related to each other than either is to a lizard or snake. They are each members of a single clade, descended from a common ancestor not shared with lizards or snakes. Under the conventions of cladism, therefore, the term 'dinosaur' is only legitimate if it includes the birds. If it does not, then it fails to contain all the descendants of a single common ancestor. If you wish to refer only to the sort of dinosaur that looks like Godzilla and is extinct, cladists would insist that you say 'non-avian dinosaur', which is ideologically pure but, if I may speak frankly, a bit clunky for use in any but a technical context. Birds, lizards, dinosaurs and snakes, of course, are all descended from a more remote common ancestor, so they all legitimately belong to a larger clade than that of the dinosaurs (and birds). Clades fit within clades, a bit like those Russian matrioshka dolls we've all seen. (BTW, cladists wince at the term 'reptile' itself, for the reasons described above. They'd let you say 'reptile' so long as you were willing to include yourself in that group. Otherwise, if you mean what we all mean when we say 'reptile', you had better say 'non-mammalian, non-avian amniote' when you are talking with a cladist.)
When the Philodromines became Philodromids, then, a lot more was going on than the mere swapping out of a couple of letters. The assertion was being made that a biggish group of animals was not a single group at all; that they did not all share a single common ancestor. (Of course the running spiders and thomisid crab spiders do share a common ancestor in any case; but that ancestor was being pushed back at least one level.)
This might all seem very rarified and academic to you, but think of an example a bit closer to home. Are you a chimpanzee? You might be tempted to answer indignantly that you are not. And, in a tightly focused context, you may legitimately distinguish yourself from chimpanzees - today's chimps and bonobos share a common ancestor that we don't have. Yet if you broaden your focus just a little a bit and wish to distinguish yourself from a gorilla, you are going to have to take our chimpanzee cousins along for the ride. For we and they descend from a shared common ancestor that arose after our lineage split from the lineage leading to gorillas. It's understandable, perhaps, that we instinctively divide the primates into two great groups - ourselves and all the others. But there is no systematically legitimate way to hive humans off from other primates unless we take the chimpanzees with us. They are no more closely related to gorillas than we are. Indeed, since the ancestors of (chimps + bonobos) split off from a common lineage after the ancestors of ((chimps + bonobos) + humans) did, from a cladistic perspective it's not so much that humans are a sort of chimpanzee as that chimpanzees are a sort of human.
Posted by Mrs Tilton at 12:01 AM | Permalink
Comments
There are the definers and the refiners, then. For one you need to be seeing the whole picture in order to understand the definition of a part. For the other the pieces can be painstakingly crafted, like delicate models, without ever contemplating that vision.
We can accept evolution, it seems to me, without going in for renaming-mania. Science isn't so much about whether the speed of light is measured in m.p.h. or km/sec but about getting one's head around where the different bits fit, and in such a way as not to exclude the view of a significant error by some future Einstein.
Records show I'm a criminal; in fact I belong to a family of idea-lovers. (All this said, those taxonomists seem to have exceptional justification in the circumstances you raise.)
Posted by: Peter at 26 Mar 2004 01:25:58
THE CAFE
The candles flicker and the
Faces gather
In the cafe of the night.
The first and the second and the third
Smile and look toward my companion:
She is pale and has scarred arms.
She looks at me in a moment's panic,
Not knowing what to say -
I squeeze her knee under the table:
Her breath calms: "Hello Miss," each say.
The pool of light surrounds
Our faces as we talk and
She forgets herself:
The first and the third have no answer -
The second shies away.
The scars have their own tale
To tell as the night wears on,
And smoke thickens - the
Words grow taut: the ideas
Concentrate.
"Hello Miss," I say, taking
Her face and kissing it.
Her anger boils, she uptips the table
Fleeing into the toilets.
The door slams shut, I smash it down;
Take my precious clown and her razor:
Seat her, love her
Among my friends.
Despair will say nothing;
Damage sees his own reflection in her; and
Death can wait.
(There are words having much in common which have their origin in very different ideas)
Posted by: Peter at 26 Mar 2004 18:48:23
Maybe I had to ask *this* question ;) "Clades fit within clades, a bit like those Russian matrioshka dolls we've all seen." - That's all fine but the system itself does not help to make the decision whether to create another tuple in the decision tree or not. So in the end, doesn't the question whether there will be another matrioshka in all but the most evident cases be down to what you call "hairsplittin" between the lumpers and the splitters, to those who dispute the fact that when something walks like a duck and talks like a duck, there's a decent chance it's actually a duck...
Posted by: Tobias at 27 Mar 2004 03:47:42
A duck by any other name would quack the same? But who else but the printers of textbooks would want lots of "Peter-brother-of-the-chimp" or "Dinosaurovitch Duck" to embody these newfangled ideas of Mr Darwin's?
Posted by: Peter at 27 Mar 2004 12:12:19
Tobias,
though I hadn't wanted to go into so much detail in the original post, the system does indeed drive the decision-making. It looks at homologies and tries to decide where they fit into the grand scheme of things. That is, the system looks at things ('characters', properly speaking) that organisms have in common, but it first eliminates things that two groups of animals have in common as a functional matter but which each arose through a separate evolutionary process from separate origins (the end results of which can be very similar indeed, thanks to convergence, whilst truly homologous things can look very different). So, a bat's wing is not homologous with a butterfly's wing, but it is homologous with a whale's flipper.
Once it has some homologous things to work on, the system tries to decide whether, in any given group of organisms, those things are ancestral or derived. Cladistic systematics considers only shared derived things as indicators of membership in a clade. (Cladistic systematics, I should also note, has an extensive and jaw-breaking jargon for all these concepts, but I will try to avoid it here.) Note that everything is relative, for what is ancestral from the perspective of one clade can be derived from the perspective of a larger clade within which the smaller fits. So: the four-legged body plan is ancestral to quadrapeds; the amniotic sac ancestral to amniotes (in common parlance, reptiles, birds and mammals); hair is ancestral to mammals; etc. Ova with an amniotic sac tell you nothing about where an organism fits within the clade of amniotes - and a fortiori nothing, if that were possible, about where it fits in among mammals - because the character is ancestral to those clades; but it tells you where the animal fits within the larger clade of quadrapeds, because within that clade the character is derived.
So, to continue with your example (and assuming that quacking is a character derived from a common ancestor that all ducks share with no other birds), then if it quacks, it is indeed a duck; meaning that you may place the animal in the duck clade (which nests within several other clades among the Aviales, which in turn nest within the Euraptoridae, which in turn nest among the dinosaurs, and so on ad pretty near infinitum, all within what we may call a metaclade of 'living things', as most biologists think life arose only once). But the fact of quacking is no help at all in figuring out where your animal fits within the duck clade.
I should note that the things considered in classifying an animal need not be morphological. For example, behaviours (as the example of quacking suggests) can also do; and these days molecular analysis is very big.
At bottom, though, there is always room for judgement, and hence for mistakes (or, shall we say, for differing tentative conclusions). If you look at most spider books, you will find the genus Cheiracanthium classed within the Clubionidae. You may know Cheiracanthium from its representative, sometimes found in Germany, the 'Dornfinger' (Cheiracanthium punctorium) - though if so, I hope you did not make its acquaintance by way of a bite, which would be a very painful way of acquiring knowledge. Clubionidae ('Sackspinnen') you have almost certainly seen; they are the straw-coloured mid-sized spiders who make those little silken cells you may have seen under a window-sill. Not too long ago a revision removed Cheiracanthium from the Clubionidae and placed them among the Miturgidae. If I recall the paper correctly (I have a copy, but it is tucked away someplace), this was on the basis of morphological characters, and I don't know whether anybody has done a DNA analysis of the animals. Such an analysis could conceivably support sticking Cheiracanthium back into the Clubionidae; or somebody might simply present a morphological argument that the genus does indeed belong among the Clubionidae despite some similarities to Miturgidae. It is an axiom of cladistic systematics that there is one and only one true 'tree of life'*; but it's another thing altogether to reconstruct that tree with absolute certainty, and decisions tend to be made on the basis of parsimony (in other words, the tree is pruned with Occam's razor, which is a mere logical tool). I imagine that the spiders themselves view this taxonomic turmoil with equanimity.
* Things can get a bit fuzzy down at the bottom of the tree, though. To some extent the 'tree' is reticulate rather than branching. Prokaryotes can swap genetic material between unrelated species (if one may legitimately use the term 'species' of prokaryotes, which some would dispute). And we eukaryotes bear evidence in our cells of the reticulate nature of early life; your mitochondria are the descendants of something a very remote ancestor once ate, or perhaps of something that once parasitised that ancestor. But for all that, the nesting image works once you get to even slightly advanced organisms - the term 'advanced' is strictly speaking illegitimate, of course, but you know what I mean.
Posted by: Mrs Tilton at 27 Mar 2004 13:18:41
Mrs. T.,
thanks for the long and interesting explanation!
>At bottom, though, there is always room for
>judgement, and hence for mistakes (or, shall we
>say, for differing tentative conclusions)
I suppose that was all I tried to say...
Tobias
Posted by: Tobias at 29 Mar 2004 04:01:05
But isn't the theory that eukarytotic cells evolved out of colonies of prokaryotic cells, just as functionally differentiated, multicellular organisms evolved out of colonies of single celled organisms, such as jellyfish, that developed "internal" symbiosis? By the way, you could use the word "complex" to avoid the connotations of "advanced".
Posted by: john c. halasz at 29 Mar 2004 07:21:19
John,
that is indeed the theory; or should I say, that's close to the theory. I haven't read anything suggesting that eukaryotes began as colonies of prokaryotes (though my not having read it is hardly evidence of its non-existence). Rather, the theory is that already-existing ancestral eukaryotes in some manner incorporated certain prokaryotes that, as time went by, became integral parts of the eukaryotic cell. The evidence for this seems quite strong, at least with respect to mitochondria and chloroplasts (I believe microbiologists even have a pretty good idea what free-living prokaryotes these are related to). I've seen mention of suggestions that the eukaryotic cell nucleus itself had a similar origin, but so far as I can tell that's not nearly as firmed up (assuming it is upfirmable) as it is for the aforementioned organelles.
Yes; 'complex', of course. I'd had a word free of the baggage of 'advanced' on the tip of my tongue but it just wouldn't make the leap; I suppose that was it. Thanks!
Posted by: Mrs Tilton at 29 Mar 2004 09:59:56





