The recent, brief foray into Shoebill territory made now a sensible time to use a few other Shoebill-based images I have here in the Tet Zoo archives. That, and I haven't been able to finish anything more substantive due to other commitments. We begin with a lateral view of a skull I once photographed - sorry about the crazy colours, once again my fantastic photographic skills have done me proud (this image is a scan of a piece of special paper featuring the image... I think it's called a photograph [thanks to Victor for providing modified versions]).
Note how robust the jugal is (the thick bar ventral to the eye socket). Also interesting is the dorsoposterior position of the bony nostril: obvious evidence that Balaeniceps has a trunk. No, just kidding, ha ha. In life, the nostril is covered by one of the keratinous plates of the bill and hence is not obvious: this is also the case in frigatebirds, pelicans, gannets and anhingas and is one of several morphological characters that have been used to indicate that these birds are close relatives (Mayr 2003) - a hypothesis that contradicts the 'more traditional' interpretation of the Shoebill as a weird member of Ciconiiformes, the group that includes storks and allies. Actually, it's not 'more traditional' at all, but it's definitely the view that's been most frequently presented in the popular and semi-technical ornithological literature (a very interesting, and not at all uncommon, phenomenon). The sharply hooked tip of the Shoebill premaxillae has also been suggested to be a shared derived character of shoebills, frigatebirds, pelicans, cormorants and gannets* (Cottam 1957, Mayr 2003) [other topologies are available, however: Hackett et al. (2008) found shoebills, hammerkops and pelicans to form a clade that is the sister-group to a heron-ibis clade; a clade containing frigatebirds, gannets, cormorants and anhingas was the sister-taxon to this pelican-hammerkop-shoebill + heron-ibis clade].
* It's present in newly hatched chicks but lost during growth.
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). Also neat is that the lacrimal and maxilla are fused, so the antorbital fenestra is reduced to a tiny aperture. I didn't only photograph this specimen, I also took time to draw part of the palate...
The most noteworthy thing about the Shoebill palate is that the palatines are completely fused along the midline. This is unusual, seen elsewhere only in such birds as toucans, hornbills and frogmouths (where the maxillae are fused along the midline as well, forming an extensive bony palate). On this occasion I was more interested in the fact that the palatines of this individual are strangely asymmetrical, with the right-side bone possessing a posteriorly projecting growth. I have no idea what this is and don't know what difference it made to the bird when it was alive. This reminds me: one day I'll publish a short article on neornithine palates. Anyone who's anyone knows that the two great groups of crown-birds - palaeognaths and neognaths - are classically distinguished on the basis of palatal anatomy, but the key differences are hardly ever explained usefully in the literature. I'll come back to this, be patient.
Finally, a masterpiece. I discovered the picture below by chance recently while clearing out a desk drawer: 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. The scene depicts animals preserved in the Jebel Qatrani Formation (deposited during the Late Eocene and Early Oligocene) of Egypt: the extinct shoebill Goliathia andrewsi is putting the smack down on the substantially larger Eremopezus eocaenus. Eremopezus (most recently analysed by Rasmussen et al. (2001)) is of uncertain affinities; a few similarities with palaeognaths have been mentioned on occasion, but (unlike a palaeognath) it seems to have been able to manipulate things with its toes, and superficial similarities with secretarybirds and.... shoebills were suggested by Rasmussen et al. (2001).
And, no, this is not meant to be a rigorously accurate bit of palaeontographic art: Goliathia is known only from an ulna (a partial shoebill tarsometatarsus from the Jebel Qatrani (Rasmussen et al. 1987) may also belong to it) and we have no real idea of what it looked like, while Eremopezus (looking here like a gastornithid) is known only from hindlimb elements and is also of unknown appearance (aaand I screwed up here in showing it lacking a hallux, whereas it definitely had one). As for the pigeon, err...
Having mentioned Goliathia, other fossil shoebill specimens are on record: the first is a partial distal end of a tarsometatarsus from the Upper Miocene of Tunisia, recognised as that of a shoebill by Rich (1972). Harrison & Walker (1982) later described a very similar fossil (also a partial distal end of a tarsometatarsus) from the Upper Miocene of Pakistan and suggested that it and the Tunisian fossil represented the same species, Paludiavis richae. If they're right, then shoebills once inhabited Asia as well as Africa. And, for the sake of google, let me note here that Balaeniceps has conventionally been given its own 'family', Balaenicipitidae, and - sometimes - its own 'order' too, Balaenicipitiformes.
Incidentally, for more cartoons depicting obscure fossil birds, check out the Talpanas article here. For more on the Shoebill, see...
And for more on other Cenozoic fossil birds, and on neornithine birds probably closely related to the Shoebill, see...
- Life-size two-dimensional condors and teratorns
- Titan-hawks and other super-raptors
- 2007: a good year for terror birds and mega-ducks
- You're not a proto-phorusrhacid, but you're still a cariamaen, and that's alright with me (ode to the Ameghinornithidae)
- Raven, the claw-handed bird, last of the phorusrhacids
- Gannets, most awesome of seabirds
- Fascinated by boobies
- March of the (small, plastic) pelicans
Refs - -
Cottam, P. A. 1957. The pelecaniform characters of the skeleton of the Shoebill Stork Balaeniceps rex. Bulletin of the British Museum (Natural History), Zoology 5, 51-72.
Hackett, S. J., Kimball, R. T., Reddy, S., Bowie, R. C. K., Braun, E. L., Braun, M. J., Cjojnowski, J. L., Cox, W. A., Han, K.-L., Harshman, J., Huddleston, C. J., Marks, B., Miglia, K. J., Moore, W. S., Sheldon, F. H., Steadman, D. W., Witt, C. C. & Yuri, T. 2008. A phylogenomic study of birds reveals their evolutionary history. Science 320, 1763-1768.
Harrison. C. J. O. & Walker, C. A. 1982. Fossil birds from the Upper Miocene of northern Pakistan. Tertiary Research 4, 53-69.
Mayr, G. (2003). The phylogenetic affinities of the Shoebill (Balaeniceps rex) Journal of Ornithology, 144 (2), 157-175 DOI: 10.1007/BF02465644
Rasmussen, D. T., Olson, S. L. & Simons, E. L. 1987. Fossil birds from the Oligocene Jebel Qatrani Formation, Fayum Province, Egypt. Smithsonian Contributions to Paleobiology 62, 1-20.
- ., Simons, E. L., Hertel, F. & Judd, A. 2001. Hindlimb of a giant terrestrial bird from the upper Eocene, Fayum, Egypt. Palaeontology 44, 325-337.
Rich, P. V., 1972. A fossil avifauna from the Upper Miocene Beglia Formation of Tunisia. Notes du Service gÃ©ologique 35, 29-66.
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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. ItÂ´s not as striking as your shoebill, but noticeable.
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.
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.
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.
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.
Awwww! I love the surprised (dismayed?) look on the Eremopezus' face!
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.
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).
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.
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.
Way before "sacrificed" and "euthanized"!
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.
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]
Claim itâs a parrot with its head at an odd angle.
You may have a point there.
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.
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]
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!
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.
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.
What do you mean?
Simply don't get too attached to scenarios. Phylogenetic analyses can test scenarios, not the other way around. :-|