You can never have too many shoebills

Cool! one of my favorite birds too... I find cranial bilateral assymetry kinda weird, considering that the rest of the animal (the outside at least) is often VERY symmetrical. I own a neotropical otter Lontra longicaudis skull; looking inside the braincase, around the base of the skull and the ear region, you can spot small diferences betweet the left and right sides. It´s not as striking as your shoebill, but noticeable.

I don't know what that hole is on the side of the rostrum: it might be a bullet hole, and I think it is, because if you look at the dorsal view of the same specimen below, a possible exit hole is apparent on the left side (there are two holes on the left side, actually, so I'm not sure what's happened here).

Was that bird 'collected' in the wild, or was it a captive specimen? If it's the latter, shooting would seem like an unlikely cause of death.

it was done in 1993, while I was at college, and comes from my brief phase of depicting poorly known fossil birds as interesting garish cartoons

In circa 1993, I had a Dougal Dixon-inspired phase of drawing hypothetical future animals (poorly).

I'm pretty sure the specimen concerned was collected from the wild.

Dartian asks: "Was that bird 'collected' in the wild?"

Ah yes, "collected" -- one of my favourite euphemisms. I'm surprised we've not seen it turn up in any action movies yet, as an alternative to "terminated with extreme prejudice", etc.

Wasn't a bit of palate broken?

I for one would agree that the more shoebill articles the better. You might branch out into pelicans and hamerkops too. Are there, for example, any morphological synapomorphies? Of course, it isn't just Hackett et al. that find this relationship. Even Sibley & Ahlquist did (though they didn't believe their own data on the hamerkop).

You have slightly confused Hackett et al., unfortunately. In Hackett et al., storks are the sister group of the whole group of gannets, herons, ibises, pelicans, etc. (I am calling that group Pelicaniformes, since it's the smallest group to include both pelicans and gannets. That leaves Ciconiiformes as storks, period. See the ToLWeb page.)

Dammit, I screwed up, but only because I wrote 'pelican + stork' instead of 'pelican + heron' (am now going to go correct, thanks for pointing this out). And, indeed, I meant to imply that many authors have supported a link between shoebills and pelicans, rather than between shoebills and storks (though this latter idea has been supported by some). Given this, I still wonder why inclusion of Balaeniceps in Ciconiiformes is portrayed as 'the status quo' in virtually all popular literature. In fact, before I started reading the technical literature I don't think I'd ever even encountered the idea that the Shoebill might be allied to pelicans. I wonder if the influence of, say, Wetmore was behind this.

Re: comment 5, yes, part of the palate is broken, but it concerns the maxillo-palatine region, not the pathological region on the right palatine.

Darren,
Shoebills grow in interest. Asymmetric skull, now! (Neatly apt for the cetacean-like-named Balaeniceps).

I wonder if there was a similar outgrowth (but cartilage-only) extending on the other side too, but one ossified ahead of the other? or whether that bone was extra and a morbid novelty. I'm just wondering, as I'm out of my depth here with no experience in such matters. More skulls needed!

Re the rostral holes, if they are indeed bullet wounds I would tend to think the smaller neater one would be the entry wound and the larger more ragged-edged one would be the exit wound, from what I've heard of such things (and from my experience of drilling or nailing through wood :-p).

Never underestimate the influence of Wetmore. He's the standard until shown otherwise, and it takes a very long time to show the popular literature anything. Besides, Balaeniceps has long legs. It must be a stork.

Then again, I still keep seeing the claim that NW vultures are storks too, and you can't pin that one on Wetmore.

And the holes are more likely from shotgun pellets than bullets. "Collecting" is such a nice word. But whatever the word, scientists need to keep on killing animals and preserving their remains. There's a nasty trend in Europe to take only blood samples and photos. I hope that doesn't spread.

Darren,

See if you like the less saturated versions at http://imgur.com/JTGk7.jpg and http://imgur.com/EvQgl.jpg . If you need assistance with similar corrections, I'd be glad to help.

Harshman, indeed...

Awwww! I love the surprised (dismayed?) look on the Eremopezus' face!

John:

Never underestimate the influence of Wetmore. He's the standard until shown otherwise, and it takes a very long time to show the popular literature anything.

Wetmore is to avian classification as Simpson is to mammalian classification, then.

Victor (comment 10): thanks indeed, I have replaced the saturated versions with your modified ones.

One thing about Shoebill is that nobody knows why it refuses to breed in captivity. There were just two broods in the last few years, and most Ciconiiformes nest quite readily.

Jerzy:

most Ciconiiformes nest quite readily

But as has been mentioned earlier in this thread, the shoebill is no longer considered to be a ciconiiform. Its most likely close relatives seem to be the pelicans - and pelicans, as it happens, don't breed particularly well in zoos either (considering how common they are in captivity). I don't know if too much should be made of that fact, however. Some birds reproduce well and others poorly (flamingos are another example that comes to mind) under captive conditions; there does not seem to be any clear phylogenetic or ecological pattern to it.

What a shame you've blogged about shoebills without talking about this excellent new paper!:

Smith, N.D. 2010. Phylogenetic analysis of Pelecaniformes (Aves) based on osteological data: implications for waterbird phylogeny and fossil calibration studies. PLoS One 5(10): e13354, 1-36. DOI: 10.1371/journal.pone.0013354

Ha! Smith's paper was published yesterday - the same day as my article - so, on this occasion, I think I can forgive myself for being unaware of it until now. Thanks for the heads-up. Looks like the paper supports the arrangement recovered by Mayr (2003) (where shoebills are sister to a clade that includes pelicans, frigatebirds, gannets, anhingas and cormorants).

@Dartian
These phylogenies are not accepted by majority of ornithological literature.

BTW, dinosaur people should deeply rethink their classification of theropods. If classification of modern birds based on osteology will turn to be false in multiple ways, then criteria for classification of non-avian theropods are also largely wrong and taxonomy is false.

The majority of ornithological literature is very old, out of date, and based on not much in the way of actual character data. On the other hand, the relationship of shoebills to pelicans and hamerkops is supported by vast quantities of molecular data. If it's your position that molecular data are worthless, then I suppose you are entitled to believe that there's still some doubt.

And how exactly do you suggest dinosaur people go about rethinking their classification? If osteology is no guide, what else is there to go on? First, osteology has worked quite well in many cases in birds. Molecules and morphology agree on most nodes (parrots, herons, piciforms, galloanserae, etc.). Second, dinosaurs have greater disparity than birds, since the latter are seriously constrained by the needs of flight, so (non-avian) dinosaurs have more easily observed, potentially useful characters.

Is it ignorant to be astonished that anyone would use beak details to classify birds? I can't imagine another tetrapod character so variable in response to environmental pressures.

First, osteology has worked quite well in many cases in birds. Molecules and morphology agree on most nodes (parrots, herons, piciforms, galloanserae, etc.)

My impression is that most supposed strong conflicts between molecular and morphological phylogenies have reduced as the data set for each increases in coverage (consider, for instance, what has happened with whale phylogeny). It's worth bearing in mind that many 'traditional'* morphological phylogenies were actually based on consideration of only a few supposed key characters, often with hypothetical pre-conceptions about evolutionary polarity.

*[rant] I really dislike the usage of the term 'traditional' in relation to systematics because, more often than not, such supposed 'tradition' simply doesn't exist. Wetmore's bird classification, for instance, only dates back in its most familiar form to the 1950s, and a number of alternative schemes were published before and since. 'Traditional' for most authors seems to mean "the scheme that I was taught in school".[/rant]

Chris, I strongly agree: as I noted somewhere above, many ideas that we regard as 'traditional' (e.g., that the Shoebill is a close relative of storks) are actually not traditional at all - that is, they have _not_ been widely supported in the technical literature but, rather, reflect the ideas of popular authors who relied on secondary sources. The idea that shoebills are stork relative (as in, members of Ciconiiformes) has been popular in the popular literature - but many ideas that are "popular in the popular literature" do not reflect the conclusions of those who work in the field.

Ah yes, "collected" -- one of my favourite euphemisms.

Way before "sacrificed" and "euthanized"!

Given this, I still wonder why inclusion of Balaeniceps in Ciconiiformes is portrayed as 'the status quo' in virtually all popular literature. In fact, before I started reading the technical literature I don't think I'd ever even encountered the idea that the Shoebill might be allied to pelicans. I wonder if the influence of, say, Wetmore was behind this.

By some miracle or other, Wetmore's (1960, right?) classification became the classification. Lesser textbook authors seem to have believed it was somehow mandated by the ICZN or something. Museum collections around the world are organized after it, as are apparently all popular books.

For decades people said Wetmore's classification was wrong in grouping the New World vultures with the raptors instead of the storks*, and for decades people didn't publish a classification that would have expressed this. Apparently they thought they weren't allowed to change "the official classification".

* As you've pointed out, that was actually pretty much right, funnily enough.

BTW, dinosaur people should deeply rethink their classification of theropods. If classification of modern birds based on osteology will turn to be false in multiple ways, then criteria for classification of non-avian theropods are also largely wrong and taxonomy is false.

I can only repeat: precladistic classifications are usually not based "on osteology", but on "cladistics with three characters" and/or scenario-based phylogenies. It took many people a long time to grasp the fact that you can use phylogenies to test evolutionary scenarios, not the other way around.

Furthermore, there is no sufficiently big phylogenetic analysis of neornithean phylogeny. Far from it. Even the opus magnum by Livezey & Zusi (2007), though it does have a nicely large sample of characters* and a large sample of extant taxa, has way too few taxa in general. There are almost no fossils in that huge matrix.

Making such a matrix with morphological data is simply so much work that it has never been done. With molecular data instead it's, like, a MSc thesis at the most. :-|

* Several hundred of them are parsimony-uninformative. Useless fluff. Boooo! But that still leaves over 2000 informative ones.

[from Darren: sorry, delayed by spam filter]

As for the pigeon, err...

Claim itâs a parrot with its head at an odd angle.
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@Jerzy:

@Dartian

BTW, dinosaur people should deeply rethink their classification of theropods. If classification of modern birds based on osteology will turn to be false in multiple ways, then criteria for classification of non-avian theropods are also largely wrong and taxonomy is false.

You may have a point there.
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@John Harshman:

And how exactly do you suggest dinosaur people go about rethinking their classification? If osteology is no guide, what else is there to go on?

You need subtlety and an open mind. With the latter you can make use of the kinds of approaches used in other sciences. If youâre subtle you can see there is more information in osteology matrices than traditionally used. Correlations for example. Whatâs currently done is a tiny fraction of what could be done.
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@Nathan Myers:

Is it ignorant to be astonished that anyone would use beak details to classify birds? I can't imagine another tetrapod character so variable in response to environmental pressures.

Try teeth.
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@Christopher Taylor:

My impression is that most supposed strong conflicts between molecular and morphological phylogenies have reduced as the data set for each increases in coverage (consider, for instance, what has happened with whale phylogeny).

That might be an exception but the newer data probably just plug huge holes in information that was originally far too sparse â and quite possibly not riddled with reversals. It doesnât alter the fact that molecules, with their simple advantage of being one or more steps removed from the characters that interact with the environment (which is why teeth and beaks are bad for large-scale topology) are more resistant to reversals/convergences in the physical projection.

Molecular data does come with its own problems. Saturation and long-branch attraction, for instance, are much more common there than in morphology, and hardly any extinct taxa can even be sampled (which is very bad because the strong influence of taxon selection on tree topologies has been thoroughly documented).

[from Darren: sorry, delayed by spam filter]

@David MarjanoviÄ

Molecular data does come with its own problems. Saturation and long-branch attraction, for instance, are much more common there than in morphology

Interesting. I'd guess that had something to do with the more 'random' nature of molecular mutations, in the sense of more independent of other influences. Randomness will presumably be the cause of long-branch attraction, which might be expected to be less when characters are 'clamped' by environmental constraints. This may well be a benefit on occasion.

The absence of fossils from molecular data is also a potential disadvantage. There are times of course when a fossil â one might mention Confuciusornis feducciai â is more trouble than it's worth. Just when people thought it was safe to accept the Conf. tribe hadn't inherited uncinate processes, that idiotic thing clouded the water.

All in all though, the results, and the theory (not that it's been done, but it's there, waiting) seem the favour the molecules. And they don't have the annoying feature of giving help when you don't need it and messing you up in difficult scenarios!

I'd guess that had something to do with the more 'random' nature of molecular mutations, in the sense of more independent of other influences. Randomness will presumably be the cause of long-branch attraction

Yes. When a stretch of DNA mutates at a certain rate, some nucleotide positions will mutate more than once if you leave it alone for too long. This can erase apomorphies, and often it does. All that's left then is small amounts of random similarity.

Just when people thought it was safe to accept the Conf. tribe hadn't inherited uncinate processes, that idiotic thing clouded the water.

I'd rather say all dinosaurs have uncinate processes, and they ossify whenever they feel like it. :-) This is not to say they're a useless character; whether they ossify does to some degree depend on selection pressure for a rigid ribcage. They're just another example that shows that all morphological characters in a data matrix for phylogenetic analysis should be weighted equally.

All in all though, the results, and the theory (not that it's been done, but it's there, waiting) seem the favour the molecules.

What do you mean?

And they don't have the annoying feature of giving help when you don't need it and messing you up in difficult scenarios!

Simply don't get too attached to scenarios. Phylogenetic analyses can test scenarios, not the other way around. :-|