As a treat, this week’s Taxon of the Week is a dinosaur. Even better, it’s a dinosaur with very small hands. Despite it’s small hands, it’s managed to cement itself amidst a pretty sizeable debate, yes ladies and and gentle, this week’s TotW is Limusaurus inextricabilis.
Limusaurus was discovered in 2009 by Xing Xu, a Chinese palaeontologist whose seemingly always discovering a new species of dinosaur (over 30 valid species to date). At around 1.7m in length (roughly the size of a large-ish dog), Limusaurus was far from the biggest and most exciting looking new dinosaur of 2009, however, it was certainly a bit weird:
- For starters it’s the first Asian ceratosaur ever described. Yes, you heard right a ceratosaur.
- For your main course, it’s herbiverous.
- To finish off with dessert, it’s got a beak.
Limusurus. Not your average ceratosaur.
True, whilst indeed not your average ceratosaur (or for that matter, theropod) such traits aren’t that weird in primarily carniverous clades. Within Theropoda, there’s many secondarily herbiverous taxa, incl. everyone’s favourite weirdo taxon, the therizinosaurs. Let’s also not forget the well known beaked herbiverous theropods, ornithomimosaurs. Oh, and did you know crocodylomorphs had a crack at beaks and herbivory (of course they did). Even though I keep harping on about its small hands, for frak’s sake, just take a look at alvarezsaurs (with even more pathetic arms than T. rex). So why exactly am I taking the time to blog about Limusaurus?
Mononykus trying, the meme that couldn’t be due to it’s preposterously pathetic forelimbs. Hastily drawn by me.
A debate as simple as 1, 2, 3
If you’re a fan of dinosaurs/palaeontology then you should be aware that birds evolved from theropod dinosaurs. News flash: this isn’t new. This notion really started to gain ground after seminal work by John Ostrom in the late 1960s/early 1970s (including that famous drawing of the ‘naked Deinonychus‘ as Richard likes to call it) who noted a lot of avian-like features in Deinonychus. On top of that, palaeontological records show multiple transitional forms between smaller theropods and birds (e.g. the ever eminent Archaeopteryx), and even some of the larger theropods show avian affinities (e.g. the cranial pneumatic sinuses found in Alioramus). Those are only a select few pieces of evidence that have made the evolutionary link between birds and theropods almost undeniable, there’s a so many more, it’s astounding (incl. inferred behaviour, such as avian sleeping positions in theropods: Xu & Norell 2004, and apparent egg brooding behaviour in Oviraptor: Norell et al. 1995). Watch the video if you need more peer reviewed proof that dinosaurs (even if they did look a bit like birds) were awesome. Try and show a little respect.
However, there’s always new discoveries that just love to get people arguing. The issue revolves around digit homology/identity. Essential, theropods (via evolution) ‘lost’ two fingers (digits) to arrive at the three-fingered hand seen in most theropods (aside from the aforementioned ‘weird taxa’, who lost more than 2). The same is true for birds. However, the debate revolves around which digits are lost, and which 3 form the fingers of the hand. In tetanurans (essentially the more advanced theropods, and by extension birds), it has long been thought that digits IV and V were lost, leaving a three-fingered hand consisting of digits I, II and III. Now, you’d expect birds, by being derived tetanurans to have this digital formula. Well that would be nice wouldn’t it. Unfortunately the question of I,II,II or II,III,IV in modern birds has been debated for a very long time. New evidence in the mid-late 1990s from developmental and genetic studies showed us that the three digits of the avian hand actually developed from digits II, III and IV. Gasp, a spanner in the works!
This new evidence was then used (rather wrongly) to attempt to oppose the hypothesis that birds evolved from dinosaurs (Feduccia 2002). While I agree, the 90s developmental evidence from modern birds creates some novel evolutionary dynamics to investigate, it cannot be used to deny the whole host of other evidence that links dinosaurs to birds. Following on from the developmental studies, genetic studies in the early 2000s showed that during the ontogeny of some birds, the digit identity would change from the initial II,III,IV to I,II,III. This led to the occurrence of the ‘Frame Shift’ hypothesis, which suggests that certain genetic pathways associated with dinosaur/avian digit identity allow for ‘rapid’ changing of digit homology throughout dinosaurian/avian evolution (at it’s core, and that’s a very simplified summary). Not to be bogged down by detail (genetics isn’t my strong suit), developmental geneticists thought they’d cracked it, and that the change in digit identity over avian/dinosaurian evolution was likely to have been caused by these frame shifts.
Feduccia’s (2002) argument in a nutshell.
Enter Limusaurus. So, finally, we come to Limusaurus’ role in all of this. In 2009, Xu et al. looked at Limusaurus’ small hands and went “you know what, that’s a reduced digit I” (NOT ACTUAL QUOTE), which makes Limusaurus’ digit identity II, III, IV. As we discovered right at the start of this article (before the I made my lack of genetics knowledge crystal clear), Limusaurus is a ceratosaur, which isn’t a tetanuran, but a more primitive theropod, meaning that the II,III,IV digit identity may well be shared by most/all tetanurans, with Limusaurus representing an intermediate of sorts. Thus Xu et al. state that the digit identity of Limusaurus is more in favour of a slower, stepwise acquisition of the digit identity seen in advanced tetanurans, and eventually birds. But, as the famous saying goes, you know what small hands means…(small gloves?)
That’s right, a big controversy.
If you’re confused (don’t worry, so am I), this may help.
The small gloves are off
The Xu et al. (2009) caused a fairly serious debate, with authors such as Vargas et al. arguing that the digit condition seen in Limusaurus is derived, and based on developmental and genetic work that they (Vargas et al.) carried out, suggest that faster genetic shifts occurred in the evolution of birds. Xu et al. quickly responded (and quoted Arthur Conan Doyle in a somewhat dramatic conclusion) and argued that the shift proposed by Vargas is not likely when the digit (and manus) morphology of fossil tetanurans is considered.
I’d sincerely like to end this post with a succinct conclusion, saying that the debate has, over the last few years been wrapped up. However, such large debates in palaeontology, due to the very nature of our field (i.e. everything we love is dead) are rarely fully resolved. This case is no exception. Researchers from Yale (Bever et al. 2011) and other top world universities have stated time and time again that the frame shift hypothesis is still viable in the context of avian evolution, and in a recent summary by Xu and Mackem, Xu is not so sure, saying neither hypothesis has evidence to topple the other. Not one for revelling in an unnecessarily depressing ending (*coughs* Firefly) I’ll leave you with this: yes, we can’t always find all the answers to big questions in palaeontology and evolution, but by creating a synergistic relationship between palaeontology and biology (genetics, evo devo etc.), future discoveries in both fields are sure to shed some light on even the biggest of debates.
If you’d like to read more on this subject, and weren’t put off by my murdering of the genetics side of things, then I’d highly recommend the aforementioned Xu & Mackem (2013) paper for a recent summary of the field (see references below, it’s in bold).
Xu, X. et al. (2009). A Jurassic ceratosaur from China helps clarify avian digit homologies. Nature 459, 940-944.
Xu, X. & Mackem, S. (2013). Tracing the Evolution of Avian Wing Digits. Current Biology 23, R538–R544 (and references therein).
Bever, G., S. et al. (2011). Finding the frame shift: digit loss, developmental variability, and the origin of the avian hand. EVOLUTION & DEVELOPMENT 13:3, 269–279.
Vargas, A.O., Wagner, G.P. & Gauthier, J.A. in Nature Proceedings (2009).
Vargas, A.O. & Wagner, G.P. Frame-shifts of digit identity in bird evolution and Cyclopamine-treated wings. Evolution & Development 11, 163-169 (2009).
Young, R. L. et al. (2011). Identity of the avian wing digits: problems resolved and unsolved. Dev Dyn. 2011 May;240(5):1042-53.