Out today: a new Turiasaurian sauropod, Mierasaurus bobyoungi, from the Early Cretaceous Cedar Mountain formation in Utah. This comes to us courtesy of a nice paper by Royo Torres et al. (2017),

Royo-Torres et al. 2017, fig. 3. The postcranial skeleton (UMNH.VP.26004) of Mierasaurus bobyoungi gen. nov, sp. nov. with the following elements: (a) middle cervical vertebra (DBGI 69 h) in right lateral view; (b) middle cervical vertebra (DBGI 69G1) in right lateral view; (c) anterior cervical vertebra (DBGI 165) in right lateral view; (d) anterior cervical vertebra (DBGI 69G2) in right lateral view; (e) atlas (DBGI 5I) in anterior view; (f) atlas (DBGI 5I) in right lateral view; (g) posterior cervical vertebra (DBGI 95) in right lateral view; (h) posterior cervical vertebra (DBGI 19 A) in right lateral view; (i) posterior cervical vertebra (DBGI 19 A) in ventral view; (j) middle cervical vertebra (DBGI 38) in right lateral view; (k) middle cervical vertebra (DBGI 38) in dorsal view; (l) middle cervical vertebra in posterior view; (m) middle cervical vertebra (DBGI 38) in left lateral view; (n) right anterior cervical rib (DBGI 5D) in medial view; (o) right anterior cervical rib (DBGI 28 A) in medial view; (p) right anterior-middle cervical rib (DBGI 95 C) in medial view; (q) right middle cervical rib (DBGI 45 F) in dorsal view; (r) right middle cervical rib (DBGI 95 A) in dorsal view; (s) left anterior cervical rib (DBGI 95B) in lateral view; (t) left middle cervical rib (DBGI 95 H) in lateral view; (u) left middle cervical rib (DBGI 95D) in dorsal view; (v) right posterior cervical rib (DBGI 10) in dorsal view. A plus sign (+) indicates a diagnostic character for Mierasaurus bobyoungi gen. et sp. nov. An asterisk (*) indicates an autapomorphy of Mierasaurus bobyoungi gen. et sp. nov. (© Fundación Conjunto Paleontológico de Teruel-Dinópolis) in Adobe Illustrator CS5 (www.adobe.com/es/products/illustrator.html).

[Because this paper is in Nature’s Scientific Reports, it inexplicably has a big chunk of manuscript chopped out of the middle, supplied separately, not formatted properly, and for all we know not peer-reviewed. This includes such minor details as the specimen numbers of the elements that make up the holotype, and the measurements. Note to self: rant about how objectively inferior Scientific Reports is to PeerJ and PLOS ONE some time.]

Anyway, this is a nice specimen represented by lots of decent material, including plenty of presacral vertebrae, which is great.

But here’s where it gets weird. Until now, Turiasauria has been an exclusively European clade. Just like Diplodocidae used to be an exclusively North American clade until Tornieria turned up, and Dicraeosauridae used to be an exclusively Gondwanan clade until Suuwassea turned out to be a dicraeosaur, and so on.

I mentioned this in an email to Matt. His initial take was:

There is a semi-tongue-in-cheek biogeography “law” that states “Everything is everywhere, and the environment selects”.

It is kinda blowing my mind that so many taxa were shared between North America, Europe, and Africa in the Late Jurassic and yet we don’t see any turiasaurs in North America until the Cretaceous. I wonder if they are there in the Morrison and just not recognized — either some of the undescribed or undiscovered northern-Morrison weirdness, or currently lumped in with Camarasaurus.

I responded “That’s one read. Another is that we’re seeing convergence on similar eco-niches within widely different clades, and our analyses are not figuring this out.”

What I mean is this: what if our “Brachiosauridae” clade is really just a collection of not-closely-related taxa in the tall-shouldered very-high-browser ecological niche? And what if our “Dicraeosauridae” clade is just a collection of short-necked grazers, with independent evolutionary origins, but all converging on morphology that suits the same lifestyle?

And that is the thought that is currently freaking me out.

Royo-Torres et al. 2107, fig. 4. The postcranial skeleton (UMNH.VP.26004) of Mierasaurus bobyoungi gen. nov, sp. nov. with the following elements: (a) anterior dorsal vertebra (DBGI 54 A) in posterior view; (b) anterior dorsal vertebra (DBGI 54 A) in anteroventral view; (c) neural arch of a middle dorsal vertebra (DBGI 37) in right anterolateral view; (d) posterior neural arch of a dorsal vertebra (DBGI 19 A) in posterior view; (e) anterior dorsal vertebra (DBGI 16) in right lateral view; (f) anterior dorsal vertebra (DBGI 16) in posterior view; (g) posterior dorsal vertebra (DBGI 16) in anterior view; (h,i) posterior dorsal vertebra (DBGI 100NA 1) in anterior view; (j,k) posterior dorsal vertebra (DBGI 100NA 1) in posterior view; (l) posterior dorsal vertebra (DBGI 100NA 1) in left lateral view; (m) middle dorsal vertebra (DBGI 11) in anterior view; (n) centrum of a posterior dorsal vertebra (DBGI 24B) in ventral view; (o) centrum of a posterior dorsal vertebra (DBGI 24B) in anterior view; (p) centrum of a posterior dorsal vertebra (DBGI 192) in ventral view; (q) anterior-middle caudal vertebra (DBGI 23B) in anterior view; (r) anterior-middle caudal vertebra (DBGI 23B) in right lateral view; (s) posterior neural arch of a posterior caudal vertebra (DBGI 48) in left lateral view; (t) posterior caudal vertebra (DBGI 21) in anterior view; (u) posterior caudal vertebra (DBGI 21) in right lateral view; (v) distal caudal vertebra (DBI 37-34-529) in right lateral view; (W) anterior caudal vertebra (DBGI 192) in posterior view. For abbreviations see supplementary information. (i), (k) and (l) were drafted by R.R.T. (© Fundación Conjunto Paleontológico de Teruel-Dinópolis) in Adobe Illustrator CS5 (www.adobe.com/es/products/illustrator.html).

When I mentioned this possibility to Matt, he shared my existential terror:

What haunts me is this: we know from mammals and extant reptiles that morphological analyses suck. Laurasian moles, African moles, and Australian moles all look the same, despite evolving from very different ancestors. Ditto wolves and thylacines, horses and litopterns, etc.

Matt reminded of a paper we’ve talked about before (Losos et al. 1998), showing that this is exactly what happens with Caribbean anole lizards. Each island has forms that live on the ground, on the trunks of trees, and on branches. Phylogenetic analyses based on morphology put all the ground-livers together, ditto for trunk-climbers, ditto for branch-climbers. But molecular analyses show that each island was colonized once and the ground, trunk, and branch forms evolved separately for each island.

What if “turiasaur”, “brachiosaur”, and “titanosaur” are the sauropod equivalents? For “Caribbean island” read “continent”; for “lizard species”, read “sauropod clade”.

Will we ever know?

Matt is hopeful that we will. He’s confident that in time, we’ll get molecular analyses of dinosaur relationships — that it’s just a matter of time and cleverness. When that happens, things could be upended bigtime.

References

 

Advertisements

Zuniceratops ontogeny - Hone et al 2016 fig 2

Various methods that may be used to determine the age/ontogenetic status of a given dinosaur specimen. Central image is a reconstruction of the skeleton of an adult ceratopsian Zuniceratops, with surrounding indications of maturity (taken from multiple sources and do not necessarily relate to this taxon). (a) Development of sociosexual signals (adult left, juvenile right—modified from [9]), (b) surface bone texture (traced from [17]), (c) large size, represented here by an ilium of the same taxon that is considerably larger than that of a known adult specimen, (d) reproductive maturity, here based on the presence of medullary bone here shown below the black arrow (traced from [18]), (e) fusion of the neurocentral arch—location of the obliterated synchondrosis indicated by black arrow (traced from [19]), (f) asymptote of growth based on multiple species indicated by black arrow (based on [20]). Central image by Julius Csotonyi, used with permission. Hone et al. (2016: fig. 2).

New paper out in Biology Letters:

Hone, D.W.E., Farke, A.A., and Wedel, M.J. 2016. Ontogeny and the fossil record: what, if anything, is an adult dinosaur? Biology Letters 2016 12 20150947; DOI: 10.1098/rsbl.2015.0947.

The idea that dinosaurs had unusual life histories is not new. The short, short version is that it is usually pretty straightforward to tell which mammals and birds are adults, because the major developmental milestones that mark adulthood – reproductive maturity, cessation of growth, macro-level skeletal fusions, histological markers of maturity – typically occur fairly close together in time. This is radically untrue for most dinosaurs, which started reproducing early, often well before they were fully grown, and for which the other signals of adulthood can be wildly inconsistent.

Puny ‘pod

We don’t talk about this much in the paper, but one aspect of dinosaur life history should be of particular interest to sauropodophiles: most of the mounted sauropod skeletons in the world’s great museums belong to animals that are demonstrably not mature. They’re not the biggest individuals – witness the XV2 specimen of Giraffatitan, the giant Oklahoma Apatosaurus, and Diplodocus hallorum (formerly “Seismosaurus”).* They’re not skeletally mature – see the unfused scapulocoracoids of FMNH P25107, the holotype of Brachiosaurus mounted in Chicago, and MB.R.2181, the lectotype of Giraffatitan mounted in Berlin. And histological sampling suggests that most recovered sauropods were still growing (Klein and Sander 2008).

* The Oklahoma Museum of Natural History does have a mounted (reconstructed) skeleton of the giant Apatosaurus, and the New Mexico Museum of Natural History has a mounted reconstructed skeleton of Diplodocus hallorum. But as nice as those museums are, in historical terms those mounts are brand new, and they have not shaped the public – and professional – conception of Apatosaurus and Diplodocus to anywhere near the same degree as the much smaller specimens mounted at Yale, AMNH, the Field Museum, and so on.

OMNH baby Apatosaurus

Apatosaurs large and small at the Sam Noble Oklahoma Museum of Natural History

Basically, very little of what we think we know about sauropods is based on animals that were fully grown – and the same problem extends to many other groups of dinosaurs.

This is kind of a methodological nightmare – a colleague on Facebook commented that he had pulled his hair out over this problem – and in the paper we suggest some ways to hopefully alleviate it. I mean, the biology is what it is, but we can minimize confusion by being really explicit about which criteria we’re using when we assign a specimen to a bin like “juvenile”, “subadult”, and so on.

Supposed Former Evolution Junkie

Personally, I’m more excited about the possibilities that dinosaur life history weirdness open up for dinosaur population dynamics and ecology.

Confession time: I am a recovering and relatively high-functioning evolutionary theory junkie. In grad school I was on the heavy stuff – I read tons of Gould and Dawkins and admired them both without being smitten by either. I took seminars on Darwin and evolutionary morphology, and lots of courses in ecology – ever mindful of Leigh Van Valen’s definition of evolution as “the control of development by ecology”. I read a fair amount of Van Valen, too, until “Energy and evolution” (Van Valen 1976) burned out most of my higher cognitive centers.

I say “recovering” evolutionary theory junkie because after grad school I mostly went clean. The problem is that dinosaurs are good for a lot of things, but exploring the inner workings of evolution is usually not one of those things. As products of evolution, and demonstrations of what is biomechanically possible, dinosaurs are awesome, and we can look at macroevolutionary patterns in, say, body size evolution or morphospace occupation, but we almost never find dinos in sufficient numbers to be able to test hypotheses about the tempo and mode of their evolution on the fine scale. I suppose I could have switched systems and worked on critters in which the machinations of selection are more visible, but for me even the charms of evolutionary theory pale next to the virulent allure of sauropods and pneumaticity.

Anyway, keeping in mind that Van Valenian dictum that evolution stands with one foot in the organism-internal realm of genes, cells, tissue interactions, and other developmental phenomena, and the other in the organism-external world of competition, predation, resource partitioning, demographics, and other ecological interactions, then it stands to reason that if dinosaurs had weird ontogenies – and they did – then they might have had weird ecologies, and weird evolution full stop. (Where by ‘weird’ I mean ‘not what we’d expect based on modern ecosystems and our own profoundly mammal-centric point of view’.)

LACM Tyrannosaurus trio - Hone et al 2016 fig 1

Three growth stages of Tyrannosaurus on display at the Natural History Museum of Los Angeles County. Hone et al. (2016: fig. 1).

Actually, we can be pretty sure that the weird ontogenies and weird ecologies of dinosaurs were intimately linked (see, for example, Varricchio 2010). Like the tyrannosaurs shown here – they didn’t all fill the same ecological niche. This casts some old arguments in a new light. Was T. rex adapted for fast running? Prrrrobably – just not as a full-size adult. The skeleton of an adult tyrannosaur is that of a 500 kg cursor pressed into service hauling around 10 tons of murder. And all of this has some pretty exciting implications for thinking about dinosaurian ecosystems. Whereas mammals tend to fill up ecospace with species, dinosaurs filled up their world with ecologically distinct growth stages.

Does all of this add up to weird evolutionary dynamics for dinosaurs? Possibly. As we say in the paper,

Correct identification of life stage also is relevant to fundamentals of evolution—if the onset of sexual reproduction substantially preceded cessation of growth in dinosaurs then the ‘adult’ phenotype may not have been the primary target of selection. In fact, once juveniles or subadults are capable of reproducing, it is conceivable a population could exist with potentially no individuals making it through the survivorship gauntlet into ‘adulthood’ and close to maximum body size. The occasional hints from the fossil record of anomalously large sauropods like Bruhathkayosaurus [51], and the Broome trackmaker [52] might be explained if many sauropods were primarily ‘subadult’ reproducers, and thus extremely large adults were actually vanishingly rare.

Did that actually happen? Beats me. But it’s consistent with what we know about sauropod life history, and with the observed scarcity of skeletally mature sauropods. And it might explain some other oddities as well, such as the high diversity of sauropods in seasonally arid environments like the Morrison Formation (see Engelmann et al. 2004), and the fact that sauropods – and large dinosaurs generally – are much larger than predicted based on the land areas available to them (see Burness et al. 2001). Because the age structure of sauropod populations was so skewed toward juveniles, the average body size of most sauropod populations was probably fairly modest, even though the maximum size was immense. So maybe a continuously reproducing population didn’t require as much food or space as we’ve previously assumed.

If we can falsify that, cool, we’ll have learned something. And if we can falsify the alternatives, that will be even cooler.

I’ll stop waving my arms now, lest I achieve powered flight and really inspire controversy. Many thanks to Dave and Andy for bringing me on board for this. It was a fun project, and we hope the paper is useful. You can read Dave’s thoughts on all of this here.

References